Relationship Between Biodiversity of Recent Pollen Spectra and Vegetation of Beech Forests in Caucasus and Carpathian Mountains*

Acta Palaeobot. 42(1): 63–92, 2002

Relationship between biodiversity of recent pollen spectra and vegetation of beech forests in Caucasus and Carpathian Mountains*

ELISO KVAVADZE1 and LEON STUCHLIK2

1 L. Davitashvili Institute of Palaeobiology, Georgian Academy of Sciences, Potochnaya Str. 4, 380004 Tbilisi, Republic of Georgia; e-mail: [email protected] 2 W. Szafer Institute of Botany, Polish Academy of Sciences, Lubicz Str. 46, 31-512 Krakow, Poland; e-mail: [email protected]

Received 9 October 2000; accepted for publication 14 May 2002

ABSTRACT. The palynological diversity of surface samples from beech forests (Fagus orientalis in the Caucasus and Fagus sylvatica in the Carpathians) is characterized and its relationships to the species composition of modern vegetation is discussed. Some problems of beech ecology, relevant for the reconstructions of palaeovege- tation are considered. The problems of long-distance transport of beech pollen as well as the pollen productivity of the two beech species in various ecological conditions are discussed.

KEY WORDS: Fagus orientalis, Fagus sylvatica, pollen spectra, Caucasus, Carpathian Mountains

INTRODUCTION

The beech genus is a typical representative occurred in the Quaternary period (Shatilova of a moderate mesophilous flora and is nowa- 1974, 1977, Shatilova & Ramishvili 1990, Cho- days widespread in the northern hemisphere chieva 1980). (Fig. 1). Most of the 10 species occur in the In the northern part of the area beech mountains within the altitudinal range 1000– species are spread essentially on the plains, 2000 m a.s.l. (Fig. 2). The origin of the genus is and in the southern part in a particular moun- East Asia, where the first primitive ancestors tain belt (Fig. 2). In Europe the northern were found, evolving to the North-American boundary of beech is limited, first of all, by low and Euro-Caucasian species (Takhtadjan 1981). winter temperatures, but the southern bound- The closest species of this genus are Fagus ary is connected with the summer humidity sylvatica L. and Fagus orientalis Lipsky. The and temperature regime. A similar situation is latter is somewhat intermediate between also observed in the mountains, where the Fagus sylvatica L. and Fagus sieboldii Engl. upper limit of beech forests is mainly control- which grows in Japan (Dolukhanov 1989). led by winter temperatures and precipitation In fossil floras beech is known in the Cauca- and the lower limit by summer precipitation. sus and Europe since the Neogene (Kolakovski In the Caucasus beech forests containing 1964, S´ rodon´ 1985). In Poland beech was wide- Fagus orientalis exhibit maximum develop- spread twice: in the Tertiary and in the Ho- ment and productivity at those altitudes locene (Ralska-Jasiewiczowa 1983, S´ rodon´ where the mean temperature of the warmest 1990). However, in the Caucasus beech forests month reaches 17–19oC, with the annual temperature amplitude is no more than 21.5°C, o * This research was supported in part by the Mianowski most frequently 18–21 C, and where the an- Fund (in 2000). nual rainfall is no less than 700 mm (Dolukha- 64

10°

60°

20°

50°

40°

10°

20°

30°

10°

40°

0°

50°

30°

140°

120°

100°

80°

80° Fig. 1 Fig.

60°

60° . Distribution area of the genus Fagus genus the of area Distribution .

40°

20°

0°

20° (after Boratyn

20°

40°

40° ´ska & Boratyn

60°

60° ski 1990, emended)´

80°

100°

120°

140°

160°

180°

10°

20°

50°

20°

70°

40°

30°

40°

50°

30°

60°

10°

0° 65

Fagus sylvatica Fagus orientalis

B Fagus sylvatica Fagus orientalis N SOUTHERN NORWAY 60o

SOUTHERN SCANDINAVIA RITSA

55o TSISKARA

BAKHMARO HARZ

50o WESTERN CARPATHIANS KHODZHAL

BAVARIAN ALPS AMTKEL SOUTHERN CARPATHIANS 45o CRIMEA ADANGE

CENTRAL APENNINES OSSETIA RHODOPE MTS 40o OLYMPUS BEVRETI

ETNA LAGODEKHI

500 1000 1500 2500 2500 m a.s.l. 500 1000 1500 2000 m a.s.l.

Fig. 2. Distribution area of Fagus sylvatica L. (after Boratyn´ska & Boratyn´ ski 1990, emended) and Fagus orientalis Lipsky (after Browicz 1982, Sokolov et al. 1977, and Dolukhanov 1989, emended). A – horizontal spreading, B – vertical spreading

nov 1989). In the mountains of West Georgia In the Carpathians the beech forest belt is this optimum is within 700–1400 m, in more situated as a whole much lower than in the humid regions of East Georgia within 1000– Caucasus. In the Ukrainian Carpathians on 1450 m, while in more arid regions within the north-eastern ridges with the cool climate, 1300–1600 m. The upper limit of beech forests the altitudinal range of Fagus sylvatica is is 2000–2100 m, almost the same in both parts 700–800 m (Stoyko & Odinak 1988). In the of the region under consideration. Beech can Polish Carpathians beech forests grow at an grow as elfin woodland only in humid climate, altitude from 850 to 1100 m. The upper limit where a long period of snow cover protects coincides with the mean annual temperature the elfin woodland against winter colds and isotherm of 4oC and July temperature of 13oC spring frosts. Under these conditions beech (Dzwonko 1990). elfin woodland ascends the mountains nearly In the Caucasus as a whole, as well as in up to the places with January isotherms -7 to Georgia, beech forests occupy half of the area -8oC and August isotherms 11–12oC (Dolukha- of all forests. In the Ukrainian Carpathians nov 1989). they account for 55% of the forest area. In the 66

High Tatra Mountains beech forests account In the Caucasus the investigations of sub- only for 2.2% (Trampler et al. 1990). fossil spectra of beech forests began much Throughout Poland beech forests occupy only later (Mamatsashvili 1972, Klopotovskaya 4.1% of all forests (Boratyn´ska & Boratyn´ski 1973, Kvavadze 1974). According to Klopotov- 1990). In Europe beech forests of Fagus syl- skaya (1973), in beech forests the maximum vatica reach their maximum values in the Bal- percentage of Fagus orientalis pollen was re- kans (up to 42% of the whole forest area). corded in the pollen spectra of moss cushions Beech forests are also well developed in (up to 64.7%). Austria, Switzerland, France, and Italy (Soko- The aim of our investigation is: 1) to estab- lov 1951). lish exact criteria for distinguishing fossil The investigation carried out by Rudolf and spectra of Caucasian and Carpathian beech Firbas in 1926 should be considered as the forests and their comparison with vegetation; very first work devoted to the study of subfos- 2) to reveal the dependence of the quantitative sil pollen spectra of mountain beech forests in content of beech pollen in the spectra in differ- Poland. In the Sudetes in the Karkonosze Mts. ent ecological conditions associated with the the above-mentioned authors studied recent vertical climatic gradient. soil samples in beech forests at an altitude from 740 to 950 m. The amount of beech pollen MATERIAL AND METHODS reached maximum values directly under beech The material was gathered during Polish-Georgian trees (up to 43%). However, even at some dis- expeditions from 1980 to 2000 (Fig. 3). Georgian beech tance from the beech forest this index dropped forests and pollen spectra formed here were studied on sharply (to 6%). 12 sites from 40 spots. In the Tatra Mountains 8 pol-

20o W 0o 20o E 40o E 60o E 60o N 60o N 0 500 km

URAL

SCANDINAVIAN MTS

50o N 50o N

A 20o W 60o E C ARPA TH I o A o 40 N N B 40 N S CAUCASUS

0o 20o E 40o E

A POLAND B Agepsta 3256 Elbrus RUSSIA 5624 7 N N 1 5 6 Guba³ówka ZAKOPANE 4 1120 Kazbek Tebulosmta 5047 4492 2 12 1 8 4073 Kominiarski Giewont 11 Tebulosmta Wierch TBILISI 1909 2 3 GEORGIA 9 10 1892 Kasprowy Wierch 1985 Kozi Wierch Œwinica Wo³owiec 2301 2291 2063 3150 Kisir Dagi TURKEY ARMENIA AZERBAIJAN 2158 SLOVAKIA 10 km 100 km Adahag Kriváò 2499 Rysy Gerlachovsky Štit 3597 2494 2655

Fig. 3. Regions of beech forest studied: A – High Tatra Mountains: 1 – Sta˛z˙yska valley, 2 – Olczyska valley; B – Caucasus Mountains: 1 – Ritsa, 2 – Tsiskara, 3 – Pontic Reservation, 4 – Khodzhal, 5 – Adange, 6 – Amtkel, 7 – Ossetia, 8 – Bevreti, 9–12 – Lagodekhi Reservation 67 len spectra from 2 sites were studied. In Ojców Na- tree limit is formed by beech elfin woodland. tional Park ( south Poland) one site in the beech forest This situation is observed in the headwaters of was studied. In some places, for comparison, soils, moss cushions, Tauber pollen trap samples were stu- the Adange, Amtkel, Kelasuri, and Big Khodz- died. The material taken in the Caucasus was treated hal rivers. However, also in Abkhazia on the in the laboratory of the Institute of Palaeobiology, southern spurs of the Greater Caucasus in the Georgian Academy of Sciences, (Tbilisi) using the region of Lake Ritsa, beech forest grows below Ertdman (1943) and Grichuk (Grichuk & Zaklinskaya the spruce-pine belt. Here in the Ritsa Reser- 1948) methods, while samples from Polish beech forests were studied in the laboratory of the W. Szafer In- vation a test site of the primary beech forests stitute of Botany, Polish Academy of Sciences (Kra- was studied. A similar site in the natural ków), using the Ertdman method (Erdtman 1943, 1960). beech forest below the belt of dark coniferous Detailed description of the subfossil spore-pollen forests was studied in Adzharia in the Tsiskari spectra taken in beech forests are made on the basis of Reservation and in Guria on the way to the the AP pollen group, taking into account also the general pollen sum of NAP. Most diagrams were Bakhmaro resort in the beech forest (Pontic plotted on the same principle. In some cases, when the Reservation). pollen spectrum of one or another sample had already been described in detail in the earlier work, we re- RITSA RESERVATION. The Ritsa Reservation peated this spectrum description (Kvavadze & Stuch- (about 16 132 ha.) situated on the southern lik 1990b, 1991, 1996). These were the spectra of Ritsa and Lagodekhi Reservations and those of beech forests slopes of the Main Caucasian Range around of Trialeti. Lake Ritsa, is mostly covered by forest vegetation. At lower altitudes (300–1200 m a.s.l.) in the forest the leading role is played by horn- RESULTS beam, lime, and box tree. At higher altitudes are pine and oriental beech. High mountain CAUCASUS. WEST GEORGIA forests consist of the Caucasian fir and beech, with a small admixture of the Sosnovski pine, In West Georgia, depending of climate hu- spruce, and birch. The description of vegeta- midity, beech forests are often situated above tion and pollen spectra in three altitudinal the dark coniferous forest belt. A similar pic- belts, lower, middle and high (Fig. 4) follows ture is mostly observed in Abkhazia, where at Kvavadze and Stuchlik (1990b). lower levels the climate is dry and unfavour- Lower belt (800 m a.s.l.) represents sample able for beech forests. In areas where beech No. 21 taken from a site about 30 km west forest grows above dark coniferous forests, the from Lake Ritsa in a beech forest with under-

Site AltitudeSample [m] numberAP NAP Spores Abies nordmannianaPicea orientalisPinus JuglansBetula regiaAlnus CarpinusCarpinusFagus caucasica orientalis orientalisQuercusCastaneaUlmusTilia sativaAcerFraxinusOstryaVacciniumSorbusIlexCorylusRhododendronEphedraHedera 20% 60 20 40 20 60 40 20 20 20 60 20 20%

1150 8

950 7

Pontic 800 6

1150

Tsiskara 4

1250

Ritsa 1200 22

800 21

Fig. 4. Subfossil spore-pollen spectra of beech forest from the Ritsa Reservation (samples 21–25); Tsiskara (samples 4–6); Pon- tic Reservation (samples 6–8) 68 growth consisting mainly of Buxus colchica sample No. 22 taken from a spruce-beech and Rhododendron mucronatum. The spore- forest. It is characterized by a high pollen and pollen spectra of the sample are characterized spore concentration and a good preservation of by arboreal pollen of 64% and herbaceous and the material. In the arboreal group, constitut- spore-bearing species of 18% each. Among ar- ing 66% of the spectrum, Abies (up to 33%), boreal plants pollen of the fir (31%) and beech Fagus (up to 16%), and Picea (up to 15%) are (21%) is prevalent. Here in the beech forest dominant. The amount of the Pinus and Alnus belt the amount of beech pollen attains its pollen is smaller, 12% and 5%, respectively. maximum values. The content of Pinus, Picea, Betula, Carpinus, Castanea, Acer, Ulmus, and Alnus pollen ranges from 8 to 12%. Pollen Tilia, and Fraxinus are represented only by a of Taxus, Quercus, Carpinus caucasica, Casta- few pollen grains. Pollen grains of shrubs (Co- nea, Tilia, Acer, and Ulmus is found in single rylus, Ilex, Rhododendron, Vaccinium, and grains. Among shrubs Ilex, Corylus, and Rho- Ephedra) are found in insignificant quantities. dodendron play a considerable role. The group The herbaceous species group is distinguished of herbaceous species is marked by a great by a large variety of plants: Apiaceae, Astera- diversity. Pollen of Apiaceae, Asteraceae, ceae, Boraginaceae, Chenopodiaceae, Faba- Ranunculaceae, Boraginaceae, and Chenopo- ceae, Geraniaceae, Plantaginaceae, Plumbagina- diaceae is most abundant, but other groups ceae, Poaceae, Polygonaceae, Ranunculaceae, (Plantaginaceae, Geraniaceae, Lamiaceae, Ar- Silenaceae, Violaceae, and others. Among temisia, and others) are found occasionally. spore-bearing species Polypodiaceae prevail, Spore-bearing species are mostly represented partially identified to the species level (Polypo- by monolete spores of Polypodiaceae (up to dium vulgare), while others (Athyrium, Dryo- 65%). The content of Hupertia selago and Bo- pteris) are identified to the generic level. The trychium lunaria spores is considerable as well. spores of Botrychium are present in a rather Among monolete spores Athyrium and Polysti- large amount. chum are identified to the level of genus. The beech-spruce forest (Tab. 1, site 2) of The forest of site 1 (sample 21) is charac- the middle belt is quite different from that of terized as a well developed more or less natu- the lower belt. It differs by lower repre- ral mountain forest with absolute dominant of sentation of Fagus orientalis in the forest Fagus orientalis covering more than 75% of (under 25% of area) and a considerable high the spot area. Other trees Carpinus caucasica amount of Picea orientalis about 25% of the Acer sonsnovskyi and A. campestre as an ad- area). The most characteristic feature of this mixture cover less than 25% of the spot area. forest is well developed undergrowth stratum Phytosociological description of the studied formed by Hedera colchica and Laurocerasus sites was carried out by the Braun-Blanquet officinalis covering more than 60% of the plot method (1951) and is presented on Table 1. area, and Ilex colchica, Fraxinus excelsior, Comparison of the pollen spectrum of Philadelphus caucasicus, Sambucus nigra, sample No. 21 with the botanical description and Ulmus sp. as admixture. The sample plot of the site shows that the content of beech pol- is situated in a rocky location and comprises len is considerable underrepresented com- trees of uneven age. pared to its actual participation in the vegeta- As one can see, the spectrum of sample No. tion cover, while the amount of fir pollen 22 reflects better than of sample No. 21 the suggests that it belongs not only to the trees existing vegetation. Picea, Fagus, and Acer, as growing on the given site but mostly blown in well as some representatives of shrubs (Fraxi- from a bordering belt. The Alnus barbara pol- nus, Ilex, and Rhododendron) are adequate len (up to 11%) is also blown in from the neigh- represented in the pollen spectrum. Abies, bouring alder stands. Buxus, Laurocerasus, Alnus, Pinus, Carpinus, Ulmus, and Tilia are Philadelphus, Rubus, and Smilax growing in overrepresented mainly by long-distance pol- the sample plot are not represented in the len from other mountain belts. spectra, probably because poor preservation of In the higher belt (1250 m a.s.l.) samples their pollen in soils. As to the herbaceous stra- Nos 23–25 from the beech-fir forest have pol- tum, only its main components are reflected in len spectra also distinguished by a high con- the spectra. centration and good preservation of pollen. Middle belt (1200 m a.s.l.) represents The pollen of arboreal species attains a maxi- 69

Table 1. Ritsa Reservation

Number of site 1 2 3 Number of site 1 2 3

Sample numbers 21 22 23–25 Herbs: Date 17.06. 17.06. 17.06. 1986 1986 1986 Asarum europaeum 2.2 – – Altitude a.s.l. in m 800 1200 1250 Fragaria vesca 2.2 + – Exposure – – NW Sanicula europaea 2.2 – 2.2 Slope degree 0 0 15 Alliaria officinalis ––+ Aristolochia sp. 1.1 – – Height of trees 20–25 25–30 30 (maximum) in m Asperula odorata 1.1 – 3.3 Diameter of tree stem 100 100 110 Borago officinalis ––+ in cm Carex sp. 1.1 – – Cover of tree layer % a 90 60 50 Cephalanthera rubra 1.1 – – Cover of shrub layer % b 30 40 5 Ichenomene indica –+1.1 Cover of herb layer % c 70 80 70 Mycelis muralis 1.1 – – Phyllitis scolopendrium 1.1 + – Trees: Salvia sp. 1.2 – – Fagus orientalis a 5.5 2.2 1.1 Symphytum grandiflorum 1.3 + 2.2 b+ 1.1+ Asplenium trichomanes +.2 1 – c+ – 1.1 Athyrium filix-femina +1.13.3 Carpinus orientalis a1.1–– Calystegia sepum +–– b+ – – Cardamine hirsuta –+1.3 c++– Carex sylvatica +–– Abies nordmanniana a+ + 3.3 Cerastium dahuricum +–– b– + + Convallaria majalis +–– Picea orientalis a– 2.2+ Dentaria bulbifera –++ b– 1.1– Dryopteris filix-mas –+3.3 Acer pseudoplatanus a– – 2.2 Epipactis latifolia +–+ b– – – Euphorbia oblongifolia ––+ c– – 1.1 Festuca gigantea –+– A. sosnovskyi a 1.1. – – Galium rotundifolium –+– b1.1– – Geranium robertianum +++ A. trautvetteri a– 1.1– Geum urbanum ––+ b+ – – Lathraea squamaria ––+ c+ – – Lathyrus niger +–– A. campestre b1.1– – Luzula pilosa –++.2 c+ – – Melica nutans ––+ Quercus iberica b++– Myosotis amoena ––1.3 Taxus baccata a– 1.1– Oxalis acetosella –++ b+ – – Paris incompleta ––+ Tilia caucasica a– 1.1– Platanthera bifolia +–– b– + – Polygonatum multiflorum –+– Ulmus sp. a – + – Polypodium vulgare –+– b– 1.1– Polystichum sp. ++– Fraxinus excelsior b– + – Prunella grandiflora +–– Shrubs: Ranunculus sp. ––+ Buxus colchica 1.1 – – Rubus sp. ––+ Rhododendron mucronatum 1.1 – – Ruscus colchica –+– Ilex colchica +2.2– Sedum stoloniferum ––+ Philadelphus caucasicus +. 1.1 – Senecio platyphylloides ––2.2 Laurocerasus officinalis +3.3– Urtica dioica ––+ Rubus caesius +–– Vaccinium arctostaphylos ––+ Sambucus nigra –+– Veratrum lobelianum ––+ Hedera colchica 2.2 4.4 – Vincetoxicum scandens ––+ Smilax excelsa 1.1 – – Viola sp. + + – 70 mum of 80%, with Abies predominant (41, 49 (up to 53%). Alder is the second dominant (up and 73%, respectively). The beech pollen takes to 20%). There are high quantities of chestnut second place (17%). Pine pollen is less signifi- (up to 13%) and pine (up to 10%). Low cant (10–15%). The pollen content of the alder amounts of spruce and fir are found. Carpinus is also small (4–5%). Then come Carpinus caucasica, Quercus, Ulmus, Tilia, Betula, and orientalis, C. caucasica, Betula, Castanea, Juglans occur even in smaller quantities. The Tilia, Acer, Vaccinium, and Rhododendron. shrub group is represented by Corylus, Hede- Herbaceous species are represented by pollen ra, Ilex, Laurocerasus, Rhododendron, Sorbus, of Poaceae, Asteraceae, Fabaceae, Campanu- and Vaccinium. The group of herbaceous la, Geraniaceae, Dipsacaceae, Polygonaceae, species is poor and contains single pollen Boraginaceae, Ranunculaceae, and others. grains of Apiaceae, Artemisia, Asteraceae, Quantitatively the Silenaceae pollen is pre- Chenopodiaceae, Poaceae, Polygonaceae and dominant. The group of spore-bearing plants Silenaceae. Spore-bearing species consist only has its own peculiarities. In this group the role of ferns. of Botrychium and Lycopodium becomes more As is evident, pollen spectra in this region essential, while that of the monolete spores of more adequately reflect the existing vegeta- ferns in the spectra decreases. tion. The role of beech in the spectrum is not The sample plot is situated in the Auadkha- reduced at all, as was observed in the spectra ra region in a well developed beech-fir forest of of beech forests of the Ritsa Reservation. The uneven age form very young to old and big transported pollen of dark coniferous species trees (Tab. 1, site 3). Comparison of the pollen growing in the upper belt is found in lower spectra with the vegetation described reveals quantities. However, here the amount of pollen that the former reflect not only the principal transported from the lower belt of chestnut- forest formations but elements of isolated as- hornbeam forests is somewhat higher (espe- sociations as well. cially Alnus ).

TSISKARA RESERVATION. At an altitude of PONTIC RESERVATION. In the beech forest 1150 m, 1 km south of the meteorological sta- belt at an altitude from 900 to 1200 m three tion near the Mtirala mountain, with the high- samples in lower, middle and upper parts on est amount of precipitation reaching up to the way from Chochkhati to the Bakhmaro re- 3000 mm a year, the test site has a well-de- sort were studied (Nos 6, 7, 8 in Fig. 4). veloped natural forest (Kvavadze & Stuchlik The first sample (No. 6) was taken at the lo- 1990a). The first stratum consists only of west level of the belt under consideration in Fagus orientalis. Its trees reach here 35m in the thicket of Quercus pontica. In the pollen height and 150 cm in diameter. However, the spectra the content of arboreal species is 78%. area of cover does not exceed 40%. The stra- In the group of herbaceous plants spore-bear- tum of shrubs is also well-developed and is ing species containing monolete spores of ferns characterized by diversity of species, among prevail. In arboreal pollen beech is predomi- which Laurocerasus officinalis, covering 50% nant (42%), and Quercus pontica, widespread of the area, is predominant. There is much Da- in the undergrowth, is the second dominant phne, Rhododendron ponticum, and Vaccinium (10%). The group of conifers consists of Abies, arctostaphylos here. Fagus, Hedera Rhododen- Picea, and Pinus pollen. They are contained in dron luteum, and Sorbus are also found. Cas- equal quantities of no more than 7–8%. There tanea sativa, Ilex colchica, Rubus, and Vibur- is little lime and Caucasian hornbeam pollen. num, are met occasionally. The stratum of Juglans regia and Alnus barbata are found in herbs is poorly developed and covers up to 5% rather small quantities. Shrubs are repre- of the area. These are mainly ferns. sented by Corylus, Sorbus, Rubus, and Va c ci- For spore-pollen analysis three soil samples nium pollen. As a whole, their amount here is were studied. The pollen spectra (Fig. 4) have not high. Among herbaceous species Apiaceae, the following peculiarities. In the common Artemisia, Asteraceae, Chenopodiaceae, Poly- group arboreal pollen prevails (91–61%). The gonaceae, and Scabiosa are met in single pol- content of herbaceous species is not high. len grains. Spore-bearing species are repre- Monolete spores of ferns are predominant. sented by Dryopteris and Pteridium spores. Among arboreal species beech pollen prevails In the vegetation on the test site under con- 71 sideration the first stratum (trees) is absent, lyx, Oxalis acetosella, Cardamine hirsuta, Me- the second stratum consists of Quercus pontica landrium album, Vaccinium arctostaphylos, shrubs and covers more than 75% of the plot Cerastium vulgatum, and others. area. The maximum shrub height reaches 3 m. In the lower part of their extent the beech Under the oak thickets a lower shrub stratum forests of Bakhmaro are reflected in the pollen composed of Laurocerasus officinalis and Rho- spectra much better than at the border with dodendron ungernii, which cover area is 100%. dark-coniferous forests ( Stuchlik & Kvavadze Ilex colchica, Vaccinium arctostaphylos and Vi- 1987, Kvavadze & Stuchlik 1988). burnum orientalis cover 35% of the surface, grow sporadically under Rhododendron and BIG KHODZHAL HEADWATERS. The river Big cherry-laurel thickets. Herbaceous species are Khodzhal is situated in the eastern part of the absent under shrub thickets. Fagus orientalis, Kodori Range and originates from the spurs of Acer trautvetteri, Sorbus aucuparia, Rubus sp., the Khodjal mount (maximum height 3313 m). and Dryopteris filix-mas are recorded only in Here forests with predominance of beech at an the marginal areas. altitude of 1170–1800 m have been studied. It In the middle part of the beech forest belt should be mentioned that in the region under sample 7 was taken. Here, unlike the above- investigation the dark-coniferous forest of mentioned test site, the first stratum is pres- spruce and fir does not form a continuous belt. ent and consists entirely of Fagus orientalis. It is discontinuous and is almost completely However, here the beech pollen content in pol- replaced by the beech forest. len spectra hardly reaches 19%. Among ar- Pollen spectra from the beech forest belt boreal species Picea orientalis is predominant. (Fig. 5, samples 2, 3, 4, 5, and 6) were studied In other respects the spectrum is the same as in three parts of the belt: lower part at alti- that of sample 6. tude 1170–1200 m a.s.l. (samples 2, 3), middle In the upper part even lover quantities of part at 1400–1500 m a.s.l. (samples 3, 4), and beech pollen are recorded in the spectrum of higher part at 1800 m a.s.l. (sample 6). sample 8, taken at the upper beech forest In the lower part beech is predominant in limit. Here it accounts only for 4%. And the the forest, but rather large quantities of chest- spruce growing in the upper belt reaches 48% nut on slopes ( sample 2) and alder in a flood- in the spectrum of sample 8. Fir pollen is the plain (sample 3) are also recorded, and Carpi- second dominant (10%). nus caucasica, Tilia caucasica, and Acer At the upper limit of beech forests the vege- trautvetteri are admixed. In the undergrowth tation has the following peculiarities. The de- Rhododendron, Vaccinium, and Ilex prevail. In scription was made on the uneven-aged forest the pollen spectrum of sample No. 2 the con- site, on the eastern slope of the Bakhmaro tent of arboreal pollen is as a maximum (84%). mountain. The tree height reaches 30m, In this group the beech and chestnut also play diameter up to 110 cm. The first stratum con- an essential role (25% each). The content of sists of beech and fir, which cover 50–75% and hornbeam and lime pollen does not exceed 9%, 25% of the area, respectively. The under- while that of alder, which does not grow near growth consists only of young specimens of the site mentioned, is as high as 14%. The Fagus, Abies and Picea, covering 75% of the amount of pollen of fir, spruce, pine, elm, oak, area. and walnut is small. Among shrubs the nut- The herbaceous stratum is taxonomically tree and Rhododendron are prevalent. The rich but represented occasionally. Beech, fir, amount of the bilberry pollen is less. Ilex is not and spruce seedlings are predominant. Here reflected in the pollen spectra at all. Among typical representatives of the herbaceous stra- herbaceous species only pollen grains of Aster- tum of beech forests (Asperula odorata, Ga- aceae are found in greater amount. Spore- lium rotundifolium, Paris incompleta, Denta- bearing plants are represented only by mono- ria bulbifera, Salvia glutinosa, Carex lete spores of Polypodiaceae (16%). sylvatica, Sanicula europaea, Dryopteris filix- In the pollen spectrum of sample 3 the fea- mas, and D. dilatata) are found. These plants tures vegetation cover is adequately reflected. are more important than those characteristic Among arboreal plants alder and beech are for spruce forests. The latter are represented predominant, constituting 34% and 18%, re- by the following species: Gentiana schistoca- spectively. The pollen content of spruce and 72

ite S AltitudeSample [m] numberAP NAP PteridophytaAbies nordmannianaPiceaPinus orientalis JuglansPterocarya regiaBetulaAlnus pterocarpaCarpinusFagus caucasica orientalisQuercusCastaneaUlmus sativaTiliaSalixVacciniumRhododendronCorylus %220 60 06020202020 20 20 60 20 20%

Ossetia 1300 5

2020 2 Amtkeli 1950 1

1950 10 Adange 1850 11

1750 12

1800 6

1500 5

Khodzhal 1400 4

1200 3

1170 2

Fig. 5. Subfossil spore-pollen spectra of beech forest along the river Big Khodzhal valley (samples 2–6); Adange river valley (samples 10–12); Amtkeli river valley, beech elfin woodland (samples 1,2); Ossetia (sample 5) pine is prevalent (up to 10%), but there is little pollen of beech (33%) and alder (27%). It pollen of fir (2%). Other arboreal plants are should be noted that in the belt of beech elfin represented by single pollen grains of lime, woodland, which forms impassable thickets, elm, and walnut. There is little pollen of Rho- beech pollen in soil does not exceed 40%, while dodendron and Corylus. in the pure beech forest it is as high as 71%. There are very few herbaceous species and Probably being suppressed, the crawling form many spore-bearing ones (56%). The abun- of arboreal plants produce much less pollen. In dance of ferns results from higher humidity of the pollen spectra of the soils under the beech soils in the flood-plains of rivers. Apart from elfin woodland one can also see pollen of long- the monolete spores of ferns, trilete spores of distance transport: Pinus – 13%, Picea – 9%, Pteridium aquilinum and Lycopodium alpi- Abies – 4%, Carpinus and Castanea in small num occur. quantities, and isolated pollen grains of Tilia, In the middle part, samples 4 and 5 taken Quercus, and Ulmus. Taxonomically the com- from pure beech forests, have pollen spectra position of herbaceous plants is extremely with up to 53–71% beech pollen, and 9–18% of diverse. Pollen of Artemisia, Boraginaceae, alder pollen. The role of hornbeam, chestnut, Plumbaginaceae, Ranunculaceae, and Silena- pine, and nut-tree is insignificant. Occasion- ceae is reflected in the spectrum as well as ally occur pollen of spruce, fir, Pterocarya, wal- that of Apiaceae, Asteraceae, Chenopodiaceae nut, elm, and bilberry. Herbaceous species are Lamiaceae, Polygonaceae, and others. rather poorly represented as single pollen Monolete ferns represent almost all the spore- grains of Apiaceae, Asteraceae, Chenopodia- bearing plants. Lycopodium alpinum occurs as ceae, Fabaceae, Poaceae, and Polygonaceae. isolated spores. Spore-bearing species (31–51%) are repre- Thanks to the absence of the dark conife- sented by the monolete spores of ferns. rous forest belt, pollen spectra reflect a real In the higher part sample 6 was taken from picture of vegetation existing here. the beech elfin woodland of the subalpine belt at an altitude 1800 m a.s.l. Its pollen spectrum HEADWATERS OF THE ADANGE RIVER VALLEY. is distinguished by the following features. The The Adange River is the right tributary of the amount of arboreal species is about 45% and of river Chkhalta and is in the region of the Ma- herbaceous species only 6%. Spore-bearing rukhi Pass, where the spurs of the Greater plants reach the highest amount (49%). Among Caucasus, Chkhalta, and the eastern margin arboreal plants the leading role is played by of Bzyb Range converge. The climate here is 73 very humid, so beech forests grow together from the border of beech elfin woodland and with dark – coniferous forests, and at the tree beech forest at an altitude 1950 m and sample limit they form the belt of pure beech forests, 2 from the very elfin woodland at an altitude changing into the beech elfin woodland. Three 2020. In sample 1 beech pollen (56%) among pollen samples, from the lower, middle, and arboreal species is predominant (Fig. 5). The upper part of the belt have been taken in vari- second dominant here is pine (16%) and alder, ous type of forest. The pollen spectra are chestnut, fir, lime, and spruce pollen is represented on Fig. 5. corded in low quantities. Birch and hornbeam The lower sample No. 12 comes from an al- are met in single grains. The group of herba- titude of 1750 m in the beech-fir forest. Here ceous plants is characterized by a rather great the admixture of spruce is very low. For spore- diversity, but as a whole their amount is not pollen spectra of arboreal species, beech pollen high. There are many spore-bearing species (43%) is prevalent. As in the vegetation, the here (46%), entirely represented by monolete subdominant is fir (19%). Single pollen grains spores of ferns. of birch, oak, willow, and hazel result from a The pollen spectrum of sample 2 is charac- small admixture of these species in this alti- terized by a slight decrease in the beech pollen tudinal belt. Pollen of other arboreal species content (41%) among arboreal species and an has been transported in low quantities from increase in the herbaceous pollen content. The the lower belts. Herbs are not numerous, only pollen content of arboreal, herbaceous, and single pollen grains of Artemisia, Asteraceae, spore-bearing species here is almost the same. Chenopodiaceae, Poaceae, and Polygonaceae In the herb group Asteraceae are predominant. are present. Many monolete spores of ferns Spore-bearing species are mainly represented (21%) are recorded, including Athyrium. by monolete spores of ferns. Lycopodium and The middle sample No. 11 comes from an al- Pteridium are recorded as single spores. titude 1850 m in the belt of pure beech forests, Thus the considered beech elfin woodland where beech pollen accounts for 69%. Pollen of spectra suggest that here, as in the head- other arboreal species is transported from long waters of Big Khodzhal and Adange, the pollen distance. The composition of herbaceous productivity of the shrub form of beech is plants is richer than in sample 12. Pollen of much lower. Apiaceae, Geraniaceae, Plumbaginaceae, Sile- naceae, and others is also found. The spore NORTH CAUCASUS contents is rather high (18%). Besides monolete spores of ferns, Botrychium lunaria OSSETIA. Soil sample 5 comes from basin of spores are present. the river Ardon in the Koban gorge at an alti- The upper sample No. 10 comes from an al- tude 1300 m in the beech forest (Fig. 5). High- titude 1950 m in the belt of beech elfin wood- mountain maple grows in low quantities land above the timberline. The pollen spec- together with beech. Arboreal pollen occurs in trum is distinguished by abundance of lower quantities than herbaceous pollen, and monolete spores of ferns (up to 50%). There spore-bearing species prevail. These are essen- are many Botrychium spores here. Among ar- tially ferns (Dryopteris, Phyllitis), amounting boreal species the transported pine pollen pre- to 40% of the whole quantity of counted sporo- vails, while beech pollen amounts only to 28%, morphs. Arboreal species and shrubs account somewhat less than in the beech elfin wood- for 35% of the spectrum. Among them the land in the Big Khodzhal valley. There is a lot transported pollen of pine (41%) prevails. The of transported pollen of Alnus here. Among second dominant is beech (up to 26%). Rather herbaceous plants Asteraceae and Silenaceae high amounts of alder, elm, and hazel pollen are predominant. Pollen of Apiaceae, Artemi- (14–9%) are recorded. Birch, hornbeam, lime, sia, Chenopodiaceae, Lamiaceae, Onagraceae, and spruce are of less importance. and others is present. In the herbaceous group pollen of ruderal plants from long-distance transport prevails. THE RIVER AMTKELI HEADWATERS. The These are mainly Asteraceae and Chenopodia- trough valley of the river Amtkeli at its head- ceae. As local elements Apiaceae, Boraginaceae, waters is separated from the Adange valley by Brassicaceae, Poaceae, and others are met. a narrow mountain strip. Sample 1 comes Pollen of Cerealia (up to 3%) is also found. 74

The spore-pollen spectra as a whole adequ- Table 2. Bevreti ately reflect regional peculiarities of the Ardon Number of site (field) 2 2a gorge, where at an altitude 1200–1600 m pine and pine-birch forests prevail. The local vege- Sample number 3,4,5 6,7,8 Date 27.05.1989 13.06.1989 tation is also characterized by the abundance Altitude a.s.l. in m 1200 1150 of ferns typical for highland beech forests and Exposure E E by the predominance of beech pollen among Slope degree 45 15 broad-leaved species. In the pine and pine- Height of trees (maximum) in m 25 15 birch forests ferns are found in very low quan- Diameter of tree stem in cm 50 50 tities both in the vegetation and in pollen spec- Cover of tree layer % a 80 100 tra (no more than 8–9%). Cover of shrub layer % b 30 10 Cover of herb layer % c 20 30 EAST TRANSCAUCASIA Trees: Fagus orientalis a3.34.4 TRIALETI RANGE. Beech-oak forests occur on b1.12.2 the extreme eastern spurs of the Trialeti c+ – Range in the neighbourhood of the village Bev- Quercus iberica a–4.4 reti, and beech-spruce forests near the village Carpinus caucasica a1.12.2 Mskhaldidi Phytosociological description of b– + the studied sites carried out by the Braun- C.orientalis a– + Blanquet method (1951) is presented after our Acer campestre a+ + previous paper (Kvavadze & Stuchlik 1991) on b–1.1 A. laetum a– + Table 2. b1.1+ Samples 3, 4, 5 were taken at an altitude of Tilia caucasica b+ – 1200 m in an uneven-aged beech-spruce forest, Ulmus sp. b + – plot 2. Description of the spore-pollen spectra Shrubs: comes from Kvavadze and Stuchlik (1991). The spore-pollen spectra are distinguished by Lonicera caucasica 1.1 + Euonymus europaeus ++ maximum content of the arboreal species (82– Rubus hirtus 1.1 + 87%). Herb content is 10–18% and spores do Crataegus sp. – + not exceed 2–3% (Fig. 6). Among trees pollen of Picea (up to 41%) and beech (up to 35%) pre- Herbs: vail. Carpinus caucasica can be considered as Asperula odorata +1.1 the third dominant (up to 19%). The participa- A. taurica –1.1 tion of Quercus and Carpinus orientalis is also Dentaria bulbifera 2.2 2.2 Geranium robertianum –+ significant, up to 7% and 6% respectively. A Sanicula europaea 2.2 – few pollen of Acer, Alnus, Betula, Juglans, and Polygonatum multiflorum ++ Tilia were found. Among shrubs only Corylus Poa nemoralis –1.1 occurs in small amounts. Among herbs adven- Primula macrocalyx –1.1 titious pollen of Artemisia and Chenopodia- Viola mirabilis ++ ceae is predominant. There is little pollen of Lathraea squamaria +– Apiaceae, Asteraceae, Carex, Plantago, Poa- Lathyrus hirsutus/niger +1.1 ceae, Polygonaceae, and Ranunculaceae. In the Luzula sylvatica +– spectrum of the soil sample taken on the bank Melica nutans 1.1 1.1 very close to the river, pollen of aquatic-palu- Anthriscus sp. – 1.1 dal vegetation was revealed (Sparganium and Fragaria vesca –+ Myriophyllum). Sporiferous species are repre- Cephalanthera rubra –+ Platanthera bifolia –+ sented by Polypodium vulgare and Dryopteris Galium aparine –+ filix-mas and monolete spores of ferns without Chaerophyllum aromaticum –+ perisporium. The pollen spectrum composition Polypodium vulgare 1.1 – compares well with that of the vegetation Dryopteris filix-mas +– growing on the experimental plot. D. carthusiana –+ The test site 2a with beech-oak forest is Serratula sp. – + situated at an altitude 1150 m. Spore-pollen Silene italica –+ 75

AP Site Altitude [m]Sample number NAP Spores AbiesPicea nordmanniana orientalisPinusJuniperusJuglansAlnusBetula regiaCarpinusCarpinusFagus caucasica orientalis orientalisQuercus UlmusCastaneaTiliaAcer sativaCorylusCerealiaCichoriumPlantagoArtemisiaChenopodiaceaePoaceaeCyperaceaeApiaceaeRanunculaceaePolygonaceaeAsterPolypodiaceae 20 20 20 20 40% 20% 60 20 20 20 20 20

1150

Bevreti 2a 6

1200

Bevreti 2 3

Fig. 6. Subfossil spore-pollen spectra of beech forests of Bevreti (samples 3–8)

spectra of samples 6, 7, and 8 taken here are tory and nature of the reservation is given in characterized by predominance of arboreal our earlier work (Kvavadze & Stuchlik 1996). plants (56–68%) The proportion of herbs is 30– We studied more comprehensively the beech 38% and that of spores 1–5% (Fig. 6). Among forests situated on the reservation territory arboreal species the following are predomi- than in other regions. Besides soil samples, nant: Fagus (up to 39%), Carpinus (up to 19%), moss cushions were investigated. In 1996 pol- and Quercus (up to 35%). Pollen of Pinus (up len monitoring began here. In the beech to 26%), which in all samples probably comes forests 5 modified Tauber traps were placed, from long-distance transport, is of adventi- where the pollen rain collected in these traps tious character. The content of the Picea pollen for each calendar year is studied (Kvavadze reaches 3–4%. Abies is represented by single 2001). Material of the last three years can be pollen grains. Acer, Alnus, Carpinus orientalis, compared with those of soils and mosses. Castanea, Juglans, Tilia, and Ulmus, are dis- At the altitudes from 690 to 2035 m and tinguished by low content of pollen. Shrubs within the forest regions with participation of are represented by Corylus and Juniperus. beech the vegetation has been recorded by the Among herbs three groups are predominant: Braun Blanquet method (1951) on 9 test sites Artemisia (up to 2,9%), Asteraceae (up to (Tab. 3). 5.2%), and Chenopodiaceae (up to 50%), There In the Lagodekhi Reservation surface sam- is some pollen of Apiaceae, Carex, Plantago, ples of soils have been studied in three alti- Poaceae, Polygonaceae, Ranunculaceae, Sile- tude forest belts: naceae, and others. Single pollen grains of The lower mountain belt — beech-chest- Sparganium are encountered. Sporiferous nut forest (690–720 m a.s.l.), site 1 and 2 taxa include Ophioglossum, Polypodium vul- (field numbers 2 and 3, Tab. 3), samples 2–8; gare and other Polypodiaceae. The middle mountain belt — beech-hornbeam forest (950 – 1650 m a.s.l.), site 3–7 LAGODEKHI RESERVATION. This reservation (field numbers 10–12 and 20–21, Tab. 3), sam- is in the extreme north-eastern part of Geor- ples 24–32 and 56–60; gia. It is one of the oldest reservations with The high mountain belt — beech-maple rich vegetation containing many Tertiary re- forest (1720–2035 m a.s.l.), site 8 and 9 (field licts and high endemism. However, here dark numbers 16 and 19, Tab. 3), samples 45–47 coniferous forests are absent. Only Pinus ha- and 52–54. mata is recorded in very low quantities. Beech The description of the vegetation and spec- forests occupy two thirds of the forest area. tra is elaborated on the basis of data taken Here optimal conditions for beech are only at from the previous work (Kvavadze & Stuchlik altitudes from 800 to 1500m (Dolukhanov 1996). The main component of the forest in all 1941). More detailed information on the his- three belts is Fagus orientalis, but pure beech 76

Table 3. Description of the beech forests in the Lagodekhi Reservation

Ordinal number 123456789 Number of site (field) (3) (4) (10) (11) (12) (21) (20) (19) (16) Sample number 2,3 4–8 24–26 27–29 30–32 58–60 55–57 52–54 45–47 Date 05.06. 05.06. 07.06. 07.06. 07.06. 09.08. 09.08. 09.06. 08.09. 1989 1989 1989 1989 1989 1989 1989 1989 1989 Altitude a.s.l. in m 690 720 950 1220 1400 1650 1675 1720 2035 Exposure SE SW S N N SE W S N Slope degree 15 30 30 45 45 15 20 45 60 Height of trees (maximum) in m303025353025303530 Diameter of tree stem (maximum) in cm 120 100 80 100 80 80 90 150 100 Cover of tree layer % a 80 90 80 90 80 90 90 80 80 Cover of shrub layer % b 40 60 30 10 20 5 30 40 70 Cover of herb layer %c30205 306080706060 Trees: Fagus orientalis a 4.4 5.5 3.3 3.3 2.2 3.3 3.3 2.2 5.5 b 3.3 3.3 3.3 1.1 2.2 1.1 2.2 3.3 4.4 c 1.1 1.1 1.1 – 1.1 – – + – Castanea sativa a1.12.2––––––– b +–––––––– c +–––––––– Carpinus caucasica a 2.2 1.1 3.3 1.1 3.3 3.3 3.3 – – b +–+––++–– c +++–––––– Fraxinus excelsior a2.21.1––––––– b +–––––––– c +–––––––– Tilia caucasica a 1.1 + 3.3 2.2 1.1 1.1 1.1 – – b ++––––1.1–– Quercus iberica/ a 1.1 + – – – 2.2 *1.1 – – *Q. macranthera b +–––––+–– c –+––––––– Acer velutinum a ––––––––– b –+––––––– c ––+1.1––––– A..campestre b1.1–––––––– A.. pseudoplatanus a ––––––––– A. trautvetteri a –––––2.22.23.31.1 b –––––––1.1– c – – – – 1.1 + 1.1 1.1 – A. laetum a – + – 1.1 1.1 – 2.2 – – b1.1+––+–++– c +––1.11.1–––– Corylus avellana b3.31.1––––+–– c ––+–––––– C. colurna a –––––1.1––– Shrubs: Rubus caesius 2.2–––––––– R. .idaeus –––––––++ R. serpens 1.1––––––++ Rubus sp. – + – 2.2 2.2 + + – + Ulmus elliptica b +–––––––– Hedera helix b +–+–––––– c ++1.1–––––– 77

Table 3. Continued

Ordinal number 123456789 Sambucus nigra b ––––+–––+ Rhododendron flavum b –1.1––––––– Sorbus cancasigena –––––––1.1 Herbs: , Asperula odorata 1.1 – + + 3.3 3.3 3.3 3.3 1.1 A. taurica –––––+1.1+1.1 Festuca sylvatica 2.2 2.2 +– – – + 1.1 2.2 – Dryopteris filix-mas + + + 3.3 1.1 3.3 3.3 3.3 3.3 D. carthusiana –––––––++ Polygonatum multiflorum +–––––+.3–+ P. verticillatum ––+––––+ Salvia glutinosa +––––––++ Viola mirabilis ++––––––– V. sylvestris ––+–++++– Euphorbia macrocarpa + – – + + 1.1 1.1 1.1 + Pteridium aquilinum +.2+––––––– Circea lutetiana ++––––––– Geranium robertianum –––––1.11.11.1– Dentaria bulbifera – – + 2.2 + + 1.1 – 1.1 Moehringia trinervia ––––+1.11.1++ Polystichum braunii –––++––—– Stellaria holostea –––––3.33.32.2– Poa nemoralis –––––+1.1–– Athyrium filix femina –––––1.1+–– Neottia nidus-avis –––+–+––– Astrantia maxima –––––++–+ Gentiana schistocalyx ––––––1.1+1.1 Carex brizoides –+––––––– C. sylvatica –––––––+– Stachys sylvatica –+––+++–– Primula macrocalyx –––––1.11.1++ Galeopsis tetrachit ––––––+–– Geum urbanum –––––++++ Phyllitis scolopendrium ––––+–––– Urtica dioica ––––+–+–– Anthriscus nemorosa –––––+–+– Calamagrostis arundinacea ––––4.4–––2.2 Impatiens noli-tangere ––––+3.33.31.3– Sedum stoloniferum –––––+++1.1 Asteraceae (cf. Telekia speciosa) –––––+1.11.11.1 Lapsana communis –––––+–++ Mycelis muralis –––––––++ Veronica chamaedrys –––––+–+– Symphytum grandiflorum –––––+–+– Senecio sp. –––––––1.1+

Sporadic species: in site nr 3: +, Aristolochia iberica +, Crataegus sp. +, Lathyrus sp. +, Salix caprea +, Vicia cracca +; 4: Carex digitata +, Hypericum androsacaeum +, Vicia crocea +; 10: Cephalantera sp.+; 12: Lathyrus sp. +; 16: Aruncus silvester 1.1, Asplenium adiantum-nigrum +, Athyrium distentifolium +, Betula litvinovii (b) +, Epilobium sp.+, Myosotis sp. +, Oxa- lis acetosella +, Ranunculus sp. +, Rumex arifolius +, Salix caprea +; 19: Cardamine hirsuta +, Galeobdolon luteum +, Ga- lium sp.+; 20: Lamium album +, Mespilus germanicus (b)+, Senecio macrophyllus 1.1, Solidago virga-aurea +; 21: Colchicum speciosum 2.2, Rumex tuberosus +, Scrophularia divaricata +, S. lateriflora +, Silene wallichiana +, Stellaria nemorum +; 21: Hesperis voronovii +. Rosa sp.+. 78 forest is very rare in the Lagodekhi Reserva- dominant. Hedera occurs as single pollen tion. Mostly these are mixed deciduous beech grains. (Fagus orientalis) forests with admixture of As already mentioned, Fagus orientalis Castanea sativa, Carpinus caucasica, Tilia dominated here in the past, and subdominants caucasica, and various Acer species. were Castanea sativa, Carpinus caucasicus, The lower mountain belt is characterized and Tilia caucasica. The undergrowth includes by considerable prevalence of Fagus oreintalis, Corylus avellana, Rhododendron flavum, which covers from 60% to more than 75% of Rubus caesius, Hypericum androsaemum, Cra- the area of sites 3 and 4 respectively, and a taegus, and Ulmus. In the herb cover Festuca significant admixture of Castanea sativa, Tilia sylvatica and Asperula odorata were predomi- caucasica, Carpinus caucasica, and Fraxinus nant. Ferns were represented by Dryopteris exelsior. Under the canopy of trees a shrub filix-mas and Pteridium aquilinum. layer with components of the tree layer and As one can see, the AP pollen spectra in Corylus avellana, Hedrea helix, Rhododendron general adequate reflect the structure of the flavum and various species of Rubus is well forest, but they show considerable underre- developed covering 40–60% of the described presentation of Fraxinus and do not show Acer site areas. In the forest studied seven soil sam- pollen in them at all. ples were taken and analysed: samples 2 and 3 In the middle mountain belt five sites of from site No. 3 and samples 4–8 from site No. 4. beech-hornbeam forest were studied (field The pollen spectra are shown on Fig. 7. The numbers 10, 11, 12, 20, 21). At lower altitude content of arboreal and shrub pollen (AP) is (950–1220 m a.s.l.) these forests grew only in high (up to 80%). The percentage of pterido- admixture with Tilia caucasica which role in phytes is 3–13%. In the AP group pollen of higher altitudes till 1675 m a.s.l. was reduced. Fagus predominates (up to 45%), and the sec- From 1220 m a.s.l. appeared Acer laetum ond subdominant is Castanea (up to 30%). which in higher altitudes at about 1650–1675 m Carpinus pollen is to 27%, and Tilia is always a.s.l. was accompanied by A. trautvetteri, present in small amounts. Sample No. 8, Quercus macranthera, and Corylus colurna. taken directly under a lime tree, is distin- All sites represent a well developed and good guished by an abundance of Tilia pollen (up to restoring forest of uneven age, with all group 45%). Pollen of Alnus, Pterocarya, Pinus and of trees from juvenile to old aged big trees up Quercus is found in small amounts. Very few to 35 m high and 100 cm stem diameter. The pollen grains of Fraxinus, Juglans, Picea, and cover degree of the tree layer in all sites is Ulmus occur, and among shrubs Corylus is rather high, varies from 80% to 90%, while

Lagodekhi Reservation

Type of forestField numberAltitude of site [m]Sample (tab. 3) numberAlnus Carpinus caucasicaCarpinusCastanea orientalis sativaCorylusFagus orientalis FraxinusHederaJuglansPicea regiaPinus orientalisPterocaryaQuercus pterocarpaTilia Ulmus Sum trees and shrubs Sum uplandSum herbs pteridophytesPollen sum

3 690

4 720

beech-chestnut 6

20% 2040 20 40 20 20 40% 20 40 60 80 20 20 100 200

Fig. 7. Subfossil spore-pollen spectra of beech-chestnut forest of the Lagodekhi Reservation (samples 2–8) 79 that of the shrub layer is rather low and varies 43%) with Tilia (up to 22%) as second domi- from 5% to 30%. nant, what is adequate to the representation From the site No. 10 samples 24, 25, and 26 in the forest. Only Carpinus as a subdominant were taken for pollen analysis. In spite of the is recorded in lower amounts than it is repre- fact that the three forest components con- sented in the forest composition. The other tained similar numbers of individuals, the pol- components of the spectra present a very in- len spectra (Fig. 8) show a considerable domin- teresting picture. Here there is very little pol- ance of Fagus (up to 50% of sum of trees and len of plants growing in the lower belts, much shrubs). Carpinus and Tilia (20% and 15% re- less than in site 10. However, the amount of spectively) were second subdominants. The AP Pinus pollen increases sharply, mainly coming group is characterized by large quantities of from the pine forests on the northern slopes of Pterocarya pterocarpa pollen (up to 20%), the Greater Caucasus in Daghestan. which has been carried here from below. This Pollen spectra (Fig. 8) of samples 30, 31, has evidently been caused by the southern ex- and 32 taken from the site 12 are distin- position of the slope and its great steepness. guished by a large Carpinus caucasica pollen Vertical slopes of the mountains serve as a content (up to 67%), followed by Fagus (up to screen where the pollen of many plants trans- 52%), and Tilia (up to 20%), what adequately ported from lower belts is trapped. In addition reflect the forest composition. Acer does not to Pterocarya, the spectra contain Alnus, Cas- appear in the spectra at all, while Acer laetum tanea, Juglans, Ulmus, and Corylus. Some pol- plays a considerable role in the forest composi- len has been carried from the higher mountain tion. The spectra do not show such shrub ele- belts too, for example, pollen of Betula and ments as Rubus, which grows extensively, Pinus. As a result of the long-distance trans- covering up to 20% of the brushwood layer (b) port from the west pollen of Picea and Abies in the forest. However, there is pollen of forest occur, while pollen grains of Artemisia, Cheno- species from neighbouring belts, namely podiaceae, and Ephedra have been brought in Alnus, Castanea, Corylus, Juglans, Pterocarya, from the steppes and forest-steppes of the low- and Ulmus. From higher altitudinal belts pol- land parts of East Georgia. There are also len of Pinus and Betula has been transported, many spores of ferns (up to 45%). while from the steppe regions of the eastern In pollen spectra of soil samples 27, 28, and and south-eastern Georgia, pollen of Artemisia 29 taken in site 11 (Fig. 8) local pollen is very and Chenopodiaceae has been brought in. The well represented. Fagus pollen prevails (up to spectra adequately register the local herbs and

Lagodekhi Reservation

Type of forestField numberAltitude of site [m]Sample (tab. 3) numberAbiesAlnus nordmannianaBetulaCarpinus caucasicaCarpinusCastaneaCorylus orientalisDaphne sativaEphedraFagus orientalisHederaJuglandPicea regiaPinus orientalisPterocaryaQuercus pterocarpaRubusSambucusTilia UlmusSum trees and shrubs Sum uplandSum pteridophytes herbs Pollen sum

10 950 25

11 1220 28

beech-hornbeam 30

12 1400 31

% 20 40 60 20 40 2040 20 20% 20 40 60 80 20 20 40 60 100200300

Fig. 8. Subfossil spore-pollen spectra of beech-hornbeam forest of the Lagodekhi Reservation (samples 24–32) 80 pteridophytes, with the amounts of the latter and spores of Botrychium lunaria, whereas fully matching the quantities in the present- single pollen grains of Abies and Picea were day communities. transported by the wind from north-western At the uppermost border of the beech-horn- Georgia. The spectra also show elements from beam forest, in both sites 21 and 20, Fagus the lower belts, including Alnus, Juglans, Pte- orientalis and Carpinus caucasica were domi- rocarya, and Ulmus. However, elements of the nants covering over 25% each of the area, and steppe region are much more poorly repre- Acer trautvetteri with Tilia caucasica subdomi- sented than on the south-facing slopes. Thus nant. As subdominants were also Quercus there is no pollen of Ephedra and very little of iberica, and Corylus colurna in site 21, and Artemisia, Cerealia, and others. There are Quercus macranthera and Acer laetum in many fern spores in the spectra. site 20. In the upper mountain belt at altitudes In pollen spectra of samples 58, 59, and 60 of 1720 and 2035 m, two sites of beech forest from the site 21 (Fig. 9) Carpinus and Fagus were studied (Nos 19 and 16), with dominance are dominants just as in the forest, and all the of Fagus orientalis and Acer trautvetteri in the subdominants are represented as well. The tree and undergrow layers, with the covering slope aspect has limited the availability of pol- degree more than 75% of the area. As sub- len from the south of the region to that of Ar- dominant in site 19 was also Acer laetum. temisia, Chenopodiaceae, Juglans regia, and Rubus idaeus and R. serpens were the only others. Pteridophytes and other local herba- additional undershrub in both sites, while in ceous plants are adequately represented in the site 16 additional Sorbus caucasigena and Be- spectra. tula litwinowii are playing a considerable role The spore-pollen spectra of samples 56 and in the forest. The herb layer in both sites was 57 from the site 20 are characterized by the very rich with Asperula odorata and Stellaria predominance of Fagus (up to 26%) and Carpi- holostea and many ferns (Dryopteris filix-mas nus (20%) pollen, and subdomince of Tilia, and D. carthusiana), as well as representatives what adequately reflect the composition of of mountainous tall forb communities (Astera- forest. High amount of Pinus pollen (19%), was ceae, Gentiana schistocalyx and others) as pre- transported from pine forests of Daghestan, dominant. since the slope is of north-western aspect here. Soil samples 52, 53 and 54 were taken from From the higher belts there was also trans- the site 19 and samples 45, 46, and 47 from ported pollen of Betula and Rhododendron, the site 16 at the upper limit of the beech-

Lagodekhi Reservation

Type of forestField numberAltitude of site [m]Sample (tab. 3) number AbiesAcer nordmannianaAlnusBetulaCarpinus caucasicaCarpinusCeltis orientalisCorylusFagus orientalisJuglansPicea regiaPinus orientalisPterocaryaQuercusRhododendron pterocarpaRubusTilia Ulmus Sum trees and shrubsSum uplandSum herbs pteridophytes Pollen sum

21 1650 59

beech-hornbeam

56 20 1675

%20 40 20 20% 204060 20 20 40 60 100 200 300 400

Fig. 9. Subfossil spore-pollen spectra of beech-hornbeam forest of the Lagodekhi Reservation (samples 56–60) 81 hornbeam forest. The spore-pollen spectra (Fig. 10). The large amount of Betula indicates (Fig. 10) of samples from the both site are the proximity of the tree limit. The spectra do characterized by a dominance of sporiferous not contain pollen of Abies and Picea, most plants (Pteridophytes) reaching their maxi- probably because of the northern exposition of mum value (62.5%). the slope. However, Pinus is always present, In pollen spectra from the site 19 among the with a high percentage of pollen (up to 10%), AP group pollen of Fagus, Pinus, and Carpinus coming mainly from long distance transport. predominates. The amount of Acer occurs only The quantities of Alnus, Corylus, and Pteroc- as single grains. Among the conifers pollen of arya are fairly significant (up to 2.7%, 4% and Picea orientalis is always present, while that 2.2% respectively), and pollen grains of Acer, of Abies nordmanniana occurs only in one Carpinus orientalis, Castanea, Ephedra, sample. The elements of the lower belts are Juglans, Quercus, Tilia, and Ulmus are com- represented by fairly significant amounts of ing from long-distance transport. Alnus barbata and Pterocarya pterocarpa. Cas- In the NAP group Polypodiaceae ( up to tanea sativa, Juglans regia, Quercus iberica, 60%) are dominants, while Chenopodiaceae Tilia caucasica, and Ulmus glabra occur only (up to 5%), Artemisia (up to 1.1%), and other sporadically. Among the shrubs pollen of Cory- Asteraceae (up to 1.1%), can be considered as lus and Ephedra is found only in two samples. subdominants, other herbaceous plants (Apia- In the NAP group Chenopodiaceae (up to ceae, Euphorbiaceae, Poaceae, Ranunculaceae, 21%), Artemisia, and Asteraceae predominate Taraxacum and others) are represented in less here on the southern slope. Among sporiferous than 1% of pollen in the spectra. There are plants, apart from Dryopteris, there are spores many spores of Dryopteris and Asplenium (up of Botrychium and Ophioglossum. The former to 12.4% and 5.5% respectively). Pollen of sy- has most probably been carried in from the nanthropic plants occurs only sporadically. upper belts, while the latter has come from Thus the factual material presented above below. Pollen of many NAP taxa is found in suggests that the maximum beech content in small amounts, included elements of synan- the spectra is recorded at an altitude 720– thropic vegetation. 950 m. The same is the case for the soil spec- Pollen spectra of samples 45, 46, and 47 tra of beech forests within the altitude range from the site 16 are characterized by domin- 1230–1400 m. However, in spite of the absence ance of Fagus (up to 36%), Carpinus (up to of the dark coniferous forest belt at its upper 18%), and Betula (up to 15%), in the AP group growth limit, beech was not adequately re-

Lagodekhi Reservation

Type of forestField numberAltitude of site [m]Sample (tab. 3) numberAbiesAcer nordmannianaAlnusBetula CarpinusCarpinus caucasicaCastaneaCorylus orientalisEphedra sativaFagus orientalisJuglansPicea regiaPinus orientalisPterocaryaQuercusRhododendron pterocarpaRubusSalixTilia UlmusSum trees andSum shrubs uplandSum herbs pteridophytes Pollen sum

19 1720 53

beech-maple

16 2035 46

% 20 40 20 20 40 20 40% 2040 20 20 40 60 100200 300 400

Fig. 10. Subfossil spore-pollen spectra of beech-maple forest of the Lagodekhi Reservation on the upper limit of Fagus orientalis (samples 52–57) 82 flected (underrepresented) in the pollen spec- maple grow. In the pollen spectra (Fig. 11) tra of soil samples. some predominance of herbaceous plants (55%) over arboreal species is characteristic. POLISH CARPATHIANS In the group of arboreal pollen, despite the predominance of beech in the forest, beech pol- In the High Tatra Mountains the beech len accounts for 6% only. In the pollen spectra forest belt (Carpathian beech forests, Dentario alder (25%) and pine (20%) pollen prevails. glandulosae-Fagetum of the Querco-Fagetea They do not grow not only on test site 14, but class) is situated at altitudes from 550 (600) m even throughout the whole beech forest belt. to 1150(1250) m a.s.l.. Besides dominating The transported birch pollen accounts for 18% Fagus sylvatica, Abies alba and Acer pseudo- of the arboreal group and should be considered platanus are occasionally admixed. as the third dominant. However, spruce, fir, Along the river Stra˛z˙yska in Tatra National and maple growing on the test site are re- Park three sites with beech forests (site 14, flected in the spectra rather poorly. The spruce 15, 16) at the altitude 800, 930, and 1000 m pollen content hardly reaches 14%, fir up to and one site along of the Olczyska valley at al- 1%, and maple no more than 1%. The content titude 870 m have been studied (more compre- of beech pollen at only 6% is also very reduced. hensive material is under study, Obidowicz Among other broad-leaved species oak, horn- personal communication). From each site beam, and elm, whose pollen was transported three soil samples were taken, while in the here from the lower belts, are also present. Olczyska valley both soil samples and moss From these belts Corylus pollen reaches rather were taken for analysis. However, not all the large quantities (up to 14%), transported from samples appeared to be suitable for analysis. long distance. Among herbaceous species only Sufficient quantities of pollen and spores could Poaceae prevails. The second dominants are be counted in 8 samples, whose spore-pollen Asteraceae species (Artemisia, Aster, Cirsium, spectra are given in Fig 11. The description and Cichorium). Quite large amounts of of the forest along the Stra˛z˙yska valley made Chenopodiaceae and other ruderal elements by the Braun Blanquet (1951) method is are found. Apiaceae, Cerealia, Plantago, and presented on Tab. 4. Ranunculaceae occur as single grains. Spore- Soil sample 31 was taken from site 14 situ- bearing species are fully represented by ated at an altitude of 1000 m, where beech and monolete spores of ferns (Dryopteris, Athy- low and equal admixtures of spruce, fir, and rium). The description of vegetation presented

Tatra Mts. – Ojców

ite S Altitude [m] Sample numberAP NAP Spores AbiesPicea alba abies Pinus sylvestrisJuniperusJuglansBetula regiaAlnus pendulaCarpinusFagus betulusQuercus sylvaticaUlmusTiliaAcerFraxinusSalixSorbus excelsiorCorylusCerealia avellanaPoaceae CyperaceaeChenopodiaceaeArtemisiaPlantagoLycopodiumPollen sum %220 40 60 0240 20 040 20 40 20 20 20 40% 200 400 600

1000 31

930 32

930 34

900 35

900 36

Str¹¿yska valley 900 37

970 11

valley 970 12

Olczyska

359 2

359 1

Ojców

Fig. 11. Subfossil spore-pollen spectra of Fagus sylvatica forest in the High Tatra Mountains: Stra˛z˙yska valley (samples 31–37); Olczyska valley (samples 11, 12); and in the Ojców National Park (samples 1, 2) 83

Table 4. Description of the beech forest in the Tatra Mts. (all records are made in the Sta˛z˙yska valley)

Number of site 14 15 16 Number of site 14 15 16

Sumple number 31 32,34 35–37 Stellaria nemorum 2.2 – – Date 13.09. 13.09. 13.09. Athyrium filix-femina 1.1 + + 1989 1989 1989 Chrysanthemum rotundifolium 1.1 – – Altitude a.s.l. in m 1000 930 900 Crepis paludosa 1.1 – – Exposure WEE Mycelis muralis 1.1 – – Slope degree 45 60 60 Phyteuma spicatum –+ + Height of trees (maximum) in m 30 25 30 Petasites albus 1.1 + + Diameter of tree stem in cm 100 50 50 Primula elatior 1.1 – – Cover of tree layer % (a) 90 100 80 Ranunculus lanuginosus 1.1 – – Cover of shrub layer % (b) 60 20 30 Actea spicata +– – Cover of herb layer % (c) 90 30 90 Ajuga reptans +1.1– Trees: Anthriscus sylvestris +– Fagus sylvatica a 4.4 5.5 4.4 Calamagrostis arundinacea ++4.4 b 1.1 1.1 2.2 Dryopteris filix-mas +– + c–++Epilobium montanum +– + Abies alba a 1.1 1.1 2.2 Fragaria vesca ++1.1 b 1.1 1.1 1.1 Luzula sylvatica +–flaves- cens+ c–++ Myosotis sylvatica +– – Picea abies a1.1–+ Gymnocarpium dryopteris +– – b2.21.1+ Polygonatum verticillatum +– – c––+ Polystichum braunii +– – Acer pseudoplatanus a1.1++ Prenanthes purpurea +2.21.1 b1.1++ Pulmonaria officinalis +– – c––+ Senecio fuchsii +– – Larix decidua a––1.1 Dentaria glandulosa –+ – Fraxinus excelsior b––+ Gentiana asclepiadea –+ + Shrubs: Hieracium lachenalii –+muro- rum 1.1 Sorbus aucuparia –+ + Lapsana communis –+1.1 Rubus hirtus –– + Mercurialis perennis –+ – R. idaeus –– + Paris quadrifolia –+ Herbs: Solidago virga-aurea –+ – Geranium robertianum 3.3 + – Streptopus amplexicaule –+ – Asarum europaeum 2.2 – + Viola reichenbachiana –1.11.1 Cardamine trifolia 2.2 2.2 – Prunella vulgaris –– + Sanicula europaea 2.2 – 1.1 Astrantia major –– + Galeobdolon luteum 1.1 – – Dryopteris carthusiana –– + Fraxinus excelsior 1.1 1.1 1.1 Soldanella carpatica –– + Veronica montana 1.1 – – Vaccinium myrtillus –– + Oxalis acetosella 3.3 2.2 3.3 Poa nemoralis –– +

in Table 4 shows that local vegetation and es- spruce (37–38%) and pine (27–28%) should be pecially participation of beech in the spectra is regarded as dominants. The third dominant is very reduced. However, the content of trans- beech (18–26%). The quantity of fir (2%) some- ported pollen is nearly 80% of the arboreal what increased. However, here the pollen con- spectrum. tent of alder and birch compared to the spectra Soil samples 32 and 34 were taken on test of sample 31 sharply decreases. In the spectra site 15, situated at altitude of 930 m. The pe- of samples 32 and 34 the oak, hornbeam, elm, culiarity of the pollen spectra is the predomin- and hazel pollen is recorded in low quantities. ance of arboreal pollen (69–67%), among which In the group of herbaceous species, like the 84 spectra of sample 31, Poaceae (27–50%) pre- quantities of transported pollen of pine (4%), vail. Rather small quantities of Asteraceae, birch (6%), and willow (1%) are found here. Chenopodiaceae and Ranunculaceae are Among broad-leaved species, besides beech, found. Among spore-bearing species ferns are oak (3%), hornbeam (0.6%), ash (0.4%), elm predominant. Lycopodium clavatum spores are (0.2%), maple (0.1%), nut-tree (1%), and lime recorded as single spores. (0.1%) are recorded, as well as hazel (2%), and For spore-pollen spectra of samples 35, 36 juniper (0.1%) of shrubs. and 37, taken at the altitude of 900 m (site The group of herbaceous pollen is also char- 16), the following features are characteristic. acterized by a rich composition, with prevail- The pollen of arboreal species (76–84%) ing of Poaceae, and large quantities of Astera- becomes more important. Among arboreal ceae, Ranunculaceae, Plantago, and Cerealia. species the transported pine pollen is predomi- Of lower importance are Cyperaceae, Chenopo- nant (39–52%). Spruce is the second dominant diaceae, and Artemisia as single grains. There (25–31%). The content of beech, as in previous are very low quantities of spore-bearing spectra, takes the third place and reaches up species, mainly monolete spores of ferns. to 11–14%. Here hornbeam, lime, and maple Pollen spectra of soil sample 12, taken from become more important. The content of fir pol- the same site of the beech forest, are charac- len as a whole also increases. However, birch, terized by the following peculiarities. The por- alder, and oak are of less importance. In the tion of arboreal species is 78% with predomin- group of herbaceous plants Poaceae also pre- ance of spruce pollen (85%). The second dominates. The importance of Cerealia and dominant is not alder, as in sample 11, but ruderal elements also increase. Among spore- pine pollen (10%). The amount of beech does bearing species ferns prevail. Athyrium and not exceed 2%, though it is the third domi- Dryopteris are identified up to the generic nant. The pollen of fir and alder is met even in level. In sample 35 Lycopodium clavatum smaller quantities. Lime and hazel are re- spores are found. corded only as single grains. The portion of On site 15 in the middle of the beech forest local arboreal pollen in the spectra of soil sam- the spectra reflected coenosis peculiarities bet- ples is 87%. ter than on site 14. Here the beech pollen con- The group of herbaceous plants is repre- tent reaches maximum values. Spruce and fir sented by abundance of Poaceae and ferns. It are also represented in higher quantities. The should be noted that in moss spectra spores of role of transported pollen is much lower than ferns are recorded in very low quantities. A in site 14 and accounts here for no more than similar regularity were observed in the ma- 35%. For site 16 the beech pollen content in terial from Lagodekhi Reservation. In general the spectra again is strongly reduced with re- Fagus is underrepresented in pollen spectra of spect to the existing vegetation, while the soil from the Tatra Mts. and pollen of conife- transported pollen of arboreal species is up rous trees is abundant thanks a high amount to 50%. of pollen from long-distance transport. While As has already been noted, along the Olczy- local pollen mainly of Poaceae is strongly over- ska valley in the High Tatra Mountains a lon- represented in the spectra. So, we should say gitudinal profile was examined from the lower that in general pollen spectra of soil in beech to the upper limit of spruce forests. In the forest of Tatra Mts. not quite adequately re- beech forest belt samples 11 (moss) and 12 flect the forest composition. (soil) at an altitude of 970 m besides beech For comparison we also studied one site in there are small quantities of fir, spruce, and the beech forest in Ojców National Park. maple and, as a whole, the vegetation charac- Beech forests here occupy nearly one third of ter is similar to that of site 14. In the spore- the forested area (Medwecka-Kornas´ & Kornas´ pollen spectrum (Fig. 11) of moss sample 11 ar- 1963). However, in the spectra of soil samples boreal species account for 64%. Transported beech-pollen content is also reduced. Pine pol- pollen of alder is predominant (35%). The sec- len is predominant (43–45%), in spite of the ond dominant is spruce (26%) and the third is fact that pine forests play a insignificant role fir (13%). The beech pollen content does not in the park. In the pollen spectra (Fig. 11) exceed 5%. Thus the portion of local pollen in among arboreal species the third dominant is moss spectra is 44%. Rather insignificant the transported pollen of birch (15%) which 85 does not grow in beech forest at all (Med- the western part of the Caucasus the total wecka-Kornas´ & Kornas´ 1992), but is common amount of arboreal species, as a whole, ex- in open landscape (Michalik 1978). Much pol- ceeds herbaceous plants considerably. len of hornbeam, alder, oak, and hazel is ob- To establish the upper tree limit in high served. Spruce, lime, elm and nut-tree are re- mountains it is very important to identify fos- corded in low quantities. sil spectra. Because in the western part of the Among herbaceous species, like the spectra Caucasus the beech elfin woodland often forms of beech forests in the Tatra Mountains, a tree line, it is necessary to identify the spec- Poaceae and Asteraceae are predominant. A lot trum of beech elfin woodland thickets in the of Chenopodiceae and Cerealia pollen is found. palaeospectrum and to distinguish it from that Apiaceae, Brassicaceae, Polygonaceae, Ranun- of the beech forest growing at lower altitudes. culaceae, Plantago, and Typha are recorded. As is evident from our material, the typical Spore-bearing species are met in single feature of the beech elfin woodland spectrum monolete spores of ferns. is the predominance of the pollen of 3 spectrum components (Tab. 5). Only the sequence of dominant and subdominant can change. DISCUSSION However, the prevailing complex is always the same and consists of Fagus-Alnus-Pinus, or The factual material presented in this work Pinus-Fagus-Alnus, or Fagus-Pinus-Alnus. High suggests that in most cases the pollen spectra content of the transported pine and alder pol- of beech forest reflect the real picture of beech len in the Caucasus conditions is charac- in the coenosis. In these spectra the beech con- teristic for open landscapes within the upper tent reaches up to 60–70%, which indicates tree limit (Klopotovskaya 1973, Kvavadze the presence of highly productive forests that 1993). Besides, for spectra being formed under are usually situated in the middle part of the thickets of beech elfin woodland the predomin- beech forest belt. However, in the marginal ance of spore-bearing species (especially ferns) areas, where the beech forest is adjacent to in the group of herbaceous plants (Tab. 5) is other vegetation belts, strong distortion of the characteristic. Noteworthy is also the nearly pollen spectra often takes place. In these cases equal ratio of arboreal pollen to spore-bearing reliable criteria to characterize the beech species. The content of other herbaceous forest spectra should be found what will allow species in the spectra considered is much one to identify similar fossil pollen spectra. lower than that of Pteridophyta repre- In the humid marine climatic conditions of sentatives. West Georgia, where the beech forest belt is As seen in Table 5, unlike elfin woodlands situated below the dark coniferous forests, ex- the spectra of beech forests have, besides tensive infestation of the spectra with conife- beech, which is almost always the spectrum rous pollen takes place. This process is most dominant, the second and third subdominants pronounced, for example, in spectra of the are usually representatives of the neighbour- Ritsa and Pontic Reservations. In this case, in ing belts, both of the lower and upper belts. In spite of spectra distortion, in all spectra of most cases this is either spruce and fir from beech forests in the group of broad-leaved above or such broad-leaved species as horn- species beech pollen still prevails (Figs 4, 5). beam, oak, and chestnut from below. It is also The same phenomenon is also observed in the important that the pollen content of beech it- spectra of beech forests of other regions of the self reaches high values (up to 60–70%) in the Caucasus. This phenomenon may be a reliable upper part as it passes into the beech elfin criterion for identification of fossil spectra of woodland. And in the beech elfin woodland beech forests due to the fact that in the Cauca- this index is no more than 28–40%. sus within the mountain forest zone beech pol- Noteworthy is also the fact that on the ter- len is not transported in large quantities from ritory of the Ritsa Reservation both in pollen one belt to another. Therefore predominance of spectra of beech-spruce and in beech-fir forests beech pollen among broad-leaved species the fir pollen is predominant (60%). Beech in means its real prevalence in the coenosis for- the beech-spruce forest is the second domi- ming the spectrum under consideration. Typical nant, while spruce, covering 25% of the arbore- is also the fact that in beech forest spectra of al stratum, occupies the third place in the 86

Table 5. The dominant pollen in beech elfin spectra and beech forest spectra in western Georgia

Beech elfin

Altitude Site name Dominant AP pollen Dominant NAP pollen Dominant forest composition

1800 m Big Khodzhal Fagus-Alnus-Pinus Polypodiaceae-Asteraceae Fagus orientalis elfin -Apiaceae – Rhododendron caucasicum 1950 m Adange Pinus-Fagus-Alnus Polypodiaceae-Silenaceae Fagus orientalis elfin -Asteraceae 1950 m Amtkel Fagus-Alnus-Pinus Polypodiaceae-Polygonaceae Fagus orientalis elfin -Asteraceae – Rhododendron caucasicum

Beech forest

1500 m Big Khodzhal Fagus-Alnus-Carpinus Polypodiaceae-Polygonaceae Fagus orientalis -Asteraceae 1750 m Adange Fagus-Pinus-Abies Polypodiaceae-Asteraceae Fagus orientalis -Polygonaceae – Picea orientalis 1600 m Kelasuri Fagus-Alnus-Abies Polypodiaceae- Asteraceae Fagus orientalis -Polygonaceae – Abies nordmanniana 1150 m Tsiskara Fagus-Alnus-Castanea Polypodiaceae- Asteraceae Fagus orientalis -Polygonaceae – Lauro-cerasus officinalis 1150 m Bakhmaro Fagus-Quercus-Picea Polypodiaceae-Asteraceae Fagus orientalis -Apiaceae – Abies nordmanniana 800 m Ritsa Reserv. Abies-Fagus-Alnus Polypodiaceae-Asteraceae Fagus orientalis-Acer -Apiaceae – Buxus – Hedera-herbs spectra. In the beech-fir forest the importance content in the spectra hardly reaches 2–4% of beech pollen is strongly reduced. However, (Kvavadze & Stuchlik 1991). However, in simi- among the broad-leaved species beech is still lar broad-leaved forests with beech participation the first dominant. The predominance of fir in the lower mountain belt of Lagodekhi the pollen in the spectra can be attributed to re- beech content in the spectra reaches 24–25%. gional peculiarities of the reservation under The peculiarity of the spectra of beech-oak consideration, where as a whole the area of fir forests with low participation of hornbeam on forests considerably predominates. However, the Trialeti Range is the predominance of on the territory of the Pontic Reservation in beech pollen (up to 39%). The second subdomi- the beech-fir forest, where beech covers 75% nant is hornbeam pollen, accounting for 19% and fir 25% of the forest area in the spectra, rather than oak, which is recorded here in the spruce pollen prevails (up to 55%), and the large quantities. The oak content in the spec- portion of fir is only 10%, while beech accounts tra accounts only for 14%, which is also ex- here for 6%. And in this case the beech pollen plained by predominance of hornbeam forests is also predominant among broad-leaved on the eastern margin of the Trialeti Range. species. High content of transported spruce Very plausible and informative are pollen pollen is also explained by a prevailing role of spectra of beech-chestnut forests in the humid spruce forests on the Bakhmaro Range. Ac- climate of the Lagodekhi Reservation. They cording to our observations, the area of fir most faithfully reflected the coenosis struc- forests here is several times smaller than that ture, where, for example, on site 4 all forest of spruce forests. dominants are represented in the pollen spec- In the conditions of East Georgia, on the tra. Beech accounts for 45%, chestnut 30%, Trialeti Range, where the climate as a whole is and hornbeam 27% of the total arboreal more arid, beech at its lower limit is more species, similar to the test site. As for beech- poorly reflected in the spectra (Fig. 6, samples hornbeam forests of the Lagodekhi Reserva- 10, 11), than, for example, in the Lagodekhi tion, they perfectly reflect the existing vegeta- Reservation with more humid climate. In tion. There beech is a dominant and hornbeam mixed broad-leaved forests with participation a subdominant. of beech on the Trialeti Range the beech pollen However, beech forests situated above 1650 m 87 are of somewhat different character. Though beech forests are more favourable. After Obi- they almost fully consist of Fagus orientalis, dowicz (1996) low quantities of beech are re- the role of the beech pollen in the spectra is corded in subfossil spectra of the Tatra Moun- strongly reduced compared to the spectra of tains. According to his data the amount of beech-hornbeam or beech-chestnut forests. beech pollen in the middle part of the beech The amount of beech pollen at an altitude forest does not exceed 34.7%, while in the mar- from 1675 to 2053 m, though being predomi- ginal areas its percentage drops to 4.7%. Not nant in the pollen spectra, hardly reaches 14– so much beech pollen is also in the spectra of 42% of the total amount of arboreal pollen. We beech forests of the Roztocze National Park in thought that this phenomenon might have south-eastern Poland, where pollen monitor- been caused by increasing importance of trans- ing is now being undertaken (Pidek & Bałaga ported pollen both from upper and lower vege- 2000). According to this author the influx of tation belts, characteristic for uplands (Kva- Fagus sylvatica pollen here is very low and ac- vadze & Stuchlik 1996). However, according to counts only for 500 gr/cm2 year. the results of pollen monitoring at the upper Researchers also report low Fagus sylvatica limit of beech forests, besides the increasing pollen productivity and its reduced role in re- the role of transported pollen, the beech pollen cent spectra, compared with other broad- influx is also very important in formation of leaved species in other European regions. The pollen spectra. At these altitudes beech pro- detailed discussion of this problem is given in duces 5–6 times less pollen than at lower le- the work by Schneider (1984), according to vels (Kvavadze 1999). which in the Kalabri Mountains in the south The comparison of peculiarities of the pollen of Italy at altitudes of 1000, 1400, and 1800 m, spectra of beech forests in the Caucasus and the beech pollen content in the beech forest Tatra Mountains showed a number of specific spectra does not exceed 14–40%. Stefanova features of recent spectra of soils being formed (1996) reported low pollen productivity of in the belt of beech forests of the High Tatra Fagus sylvatica at altitudes of 1640 and 1740 Mountains. While in the Caucasian spectra a m. According to Stefanova, in the Northern distortion of actual spectrum content takes Pirin Mountains in Bulgaria recent spectra of place essentially with the arboreal pollen moss cushions in the pure beech forest the transport from the adjacent belts, in the spec- amount of beech pollen in the spectrum hardly tra of beech forests of the Tatra Mountains reaches 21%. In this case the forest under in- large quantities of arboreal pollen from the vestigation is at the upper limit of beech. whole region are found. In no one studied pol- It should be mentioned that at the same len spectrum in the beech forest spectrum is time in the literature there are quite different beech the first dominant, since its quantity data. For example according to Arap (1984), in does not exceed 18–26%. It should be noted the belt of beech forests of the Ukrainian Car- that this maximum is observed exactly in the pathians on the Polonin Range, the beech con- spectra of the middle part of the beech forest tent of recent spectra reaches 78–84% of the belts, where, as has already been mentioned, a total arboreal pollen. A lot of beech pollen (up similar index in the Caucasian mountains can to 60%) is also recorded in other regions of the reach 60–70%. As for the marginal areas of the Ukrainian Carpathians (Arap 1984). Big quan- beech belt, in the Tatra Mountains the role of tities of beech pollen are observed in the South beech pollen is so much reduced that its pollen Carpathians. In the moss pollen spectra in the cannot even be regarded as the third subdomi- belt of beech forests of the Retezat National nant. It is evident from the quantitative beech Park in Romania at an altitude of 1110–1200 content in spectrum of sample 31 at an alti- m the beech pollen content reaches 70–90% tude of 1000 m a.s.l. in the Stra˛z˙yska valley. (Pop 1967). Here the beech content in the spectrum is 7%. Rather different results on the influx of Even smaller amounts of beech pollen (2–5%) beech pollen have been obtained in the studies are in spectra 11, and 12, obtained at an alti- of pollen rain from pollen traps and moss cu- tude of 970 m along the Olczyska valley, where shions. For example, productivity of Fagus beech reaches its upper limit, as in the orientalis is twice as low as that of Fraxinus, Stra˛z˙yska valley. Little beech pollen is also in Tilia, and Quercus and is from 5600 to 6900 Ojców National Park, where conditions for grains/cm2/year in mosses (Andersen 1970) 88 and from 8500 to 11500 grains/cm2/year in pol- addition, in these spectra the pollen of those len traps (Andersen 1974). As has already plants was recorded that are rather rarely met been mentioned, the beech pollen influx in here in the soil spectra (Acer, Juniperus, Sor- south-eastern Poland amounts to only 500 bus, and Salix). However, it should also be grains/cm2/year. However, the data on pollen mentioned that moss cushion spectra do not productivity of various arboreal species in the adequately reflect, for example, the role of Caucasus Mountains show that, on the con- spore-bearing species and especially monolete trary, Fagus orientalis produces the highest spores of ferns that are abundant in the quantity of pollen among all broad-leaved coenosis. And in soil spectra spore-bearing plants. Its maximum pollen influx at an altitude species are properly reflected. A similar situ- of 1250 and 1500 m is up to 120 000–130 000 ation is also observed from the comparison of grains/cm2/year. For comparison it should be soil and moss spectra in the Lagodekhi Reser- mentioned that in the mountains of the vation. As to the comparison of the pollen con- Lagodekhi Reservation the maximum influx of tent of beech itself in soil, moss, and pollen- Quercus iberica pollen is 83 000 grains/cm2/year trap spectra, the following regularities are Tilia caucasica up to 60 000 grains/cm2/year, observed. As a whole, in soil samples there is and Carpinus caucasica 45 000 grains/cm2/year less beech pollen than in mosses and pollen (Kvavadze 1999, 2000a). Thus, according to traps (Fig. 12). This phenomenon is quite ex- our data Fagus orientalis produces twice as plicable, since in soils, under influence of biol- much pollen as lime and oak and three times ogical and various mechanical and chemical as much as Carpinus caucasica. As has al- processes, fossilisation and the following de- ready been mentioned, at its upper limit the struction of pollen grains begins as soon as beech pollen productivity significantly de- pollen enters the soil (Tyuremnov & Berezina creases, and in the mountains of the Lagode- 1965). However, in moss cushions and pollen khi Reservation at an altitude of 2000 m it traps, which have ideal conditions due to the reaches no more than 20 000 grains/cm2/ year. absence of mineral basis and other biotic and The comparison of pollen spectra of soil abiotic factors, pollen is well conserved, and samples and moss cushions under the canopy the process of fossilisation here does not take of beech forests both in the Caucasus and place. Therefore these traps have much pollen Tatra Mountains showed that the pollen spec- of plants with thinner exine (Acer, Juniperus, trum of moss cushions is richer in its taxo- Larix, Populus, etc.) which is easily destroyed nomic composition. In mosses the total content in soils. of pollen is represented. For example, in the Analysis of our data suggests that the pol- moss spectra of sample 11 (Fig. 11) from the len spectrum of the beech forest can be mainly Olczyska valley in Tatra Mts. the amount of distorted for two reasons: 1) inadequate eco- counted pollen is much higher than in soils. In logical conditions for highly-productive beech

Lagodekhi Reservation

Fagus

in soil in moss in pollen traps

ltitude [m] A Fagus Fagus Fagus Pollen influx of 2 %2010 3040 50 10 20 30 40 50 60 10 20 30 40 50% 40000 120000 grains/cm /year

2000

1800

1500

1250

1000

500

Fig. 12. Comparison of recent spore-pollen spectra of soils and moss cushions with pollen spectra from Tauber pollen traps in beech forest of the Lagodekhi Reservation 89 forests, leading to a decrease in the pollen pro- spruce, birch, and alder is found. This phe- ductivity; 2) an increase in the amount of nomenon is also explained by the regime of transported pollen of other arboreal species local winds, whose force and duration reach both from the neighbouring mountain belts significantly higher values than in the Caucasus and from more distant regions, thus changing (Hess 1965, Lominadze & Chirakadze 1971). drastically the character of the beech-forest As for pollen spread of beech itself, both in spectrum. conditions of the Tatra Mountains and in the The simultaneous action of the above-men- studied regions of the Caucasus beech pollen tioned two factors can explain the strong dis- in big quantities cannot be transported from tortion of beech forest spectra in the Tatra one belt to another. Beech pollen also does not Mountains. Here the beech forests, compared distort pollen spectra of open landscapes of to other regions of the Carpathians, have the subalpine and alpine meadows. However, the least extent. The area of beech forests in the situation changes drastically in the Ukrainian Tatra Mountains is only 2.2%, while, for Carpathians, where most ridges have meri- example, in Bieszczady Mts. their area dional position towards west winds which reaches 40.9% and in the Beskides 27% (Tram- have high force and duration, and the area of pler et al. 1990). Such an insignificant devel- beech forests is half of the forest area. Intens- opment of beech forests in the Tatra Moun- ive beech transport begins already in spruce tains might be explained by the absence of forests, and in the alpine spectra the beech optimal conditions for growth of highly produc- content can reach up to 70% of arboreal pollen. tive forests with high pollen productivity. Our analysis showed that the actual role of beech pollen in the Tatra Mountains spectra is also CONCLUSION reduced due to mass pollen transport from other regions. The process of pollen transport The detailed analysis of regularities of for- from one vegetation belt to another by strong mation of pollen spectra of two types of beech winds in the Carpathians is described in other forests in various geographical and ecological works (Kvavadze 1988, 2000b, Stuchlik & Kva- conditions of the Caucasus and Carpathians vadze 1995). It should be mentioned here that allows us to draw a number of important con- very complex circulation of air masses not only clusions for each region, and they should be in the Carpathians but throughout the whole taken into account in interpretation of fossil territory of Poland, where during the year spore-pollen spectra. quite different air masses prevail, facilitating 1. In conditions of humid marine climate of the formation of rather peculiar and complex West Transcaucasia, in pure beech forests wind regimes. Only by winds of different direc- situated in the middle part of the beech belt, tions, including north can one explain, for adequate pollen spectra are being formed, example, too high a percentage of pine pollen where the beech content can reach 60–70%. In in the Tatra Mountains spectra (Stuchlik these spectra the second and third subdomi- & Kvavadze 1995), since the area of pine nants are usually representatives of the neigh- forests in the belts of beech or spruce forests bouring vegetation belts. on the territory of the Tatra Mountains and 2. In those spectra, where the beech role is West Carpathians does not exceed 17%. How- very reduced, the criterion for establishing a ever, in the pollen spectra the content of pine beech pollen spectrum is the predominance of pollen in the belt of beech and spruce forests beech pollen in the group of other broad-leaved can reach up to 50%, and in alpine and subal- species. In the same spectrum the total pine belts up to 60% and higher. Of course amount of arboreal pollen prevails over herba- here we deal with long-distance transport of ceous species among which only repre- pine pollen from the plains of Poland, where sentatives of Pteridophyta can reach high the area of pine forests accounts for 71.6% of values. the whole forest area. 3. The criterion for establishing the spec- In the Tatra Mountains the local pollen trum of beech elfin woodland is the presence of transport from one belt to another is higher the following three dominating components in than in the Caucasus. In the belt of beech different ratios in the pollen spectrum com- forests, apart from pine, much pollen of plex. These are either Fagus-Alnus-Pinus, or 90

Pinus-Fagus-Alnus, or Fagus-Pinus-Alnus. 9. A very low content of Fagus sylvatica pol- The pollen content of beech itself in soils len in the belt of beech forests in the High under the elfin woodland thickets does not ex- Tatra Mountain can be explained by two rea- ceed 28–40%. In the group of herbaceous sons. These are the absence of optimal condi- plants spore-bearing species and especially tions for the development of highly productive ferns are predominant. It should also be men- forests producing high quantities of pollen, tioned that the total content of fern spores ap- and intensive transport of arboreal pollen both proaches quantitatively the total amount of ar- from lower belts and from more distant re- boreal pollen. gions. 4. At the lower limit of forests with beech 10. Unlike the Ukrainian Carpathians, both participation in conditions of the more arid cli- in the Tatra and the Caucasus, beech pollen is mate of East Georgia (eastern margin of the not transported in large quantities to the Trialeti Range) the percentage of beech pollen neighbouring vegetation belts. It does not also in spectra is lower (2–4%) than in the regions distort pollen spectra of subalpine and alpine of East Georgia with the more humid climate meadows. where, for example, in the low-mountain 11. The high content of beech pollen in the forests even with the insignificant admixture fossil pollen spectra indicates the existence of of beech in the Lagodekhi Reservation its con- optimal climatic conditions for beech forests. tent reaches 24–25%. In Transcaucasia it means that temperatures 5. Among spectra of mixed beech – broad- in the warm season reached 17–19o, and in the leaved forests most adequate are pollen spec- Tatra Mountains this index is at least 13–15o. tra of beech-chestnut and beech-hornbeam Thus the character of the quantitative partici- forests of the Lagodekhi Reservation, where pation of beech pollen in the spectrum can also each coenosis component is precisely reflected be a reliable criteria for paleoclimatic recon- in pollen spectra of soil samples. In these spec- struction. tra the beech pollen content is 40–50%. 6. Pollen monitoring showed that at the upper limit of pure beech forests in cold cli- REFERENCES matic conditions of the Lagodekhi Reservation, Fagus orientalis produces 6–7 times less pol- ANDERSEN S.T. 1970. The relative pollen productiv- len than in the middle of the belt at an alti- ity and pollen representation of North European trees, and correlation factors for tree pollen spec- tude 1400–1580 m. The percentage of beech tra. Danm.Geol. Unders. II, 96: 1–99. pollen in the spectra above 1800 m hardly ANDERSEN S.T. 1974. Wind conditions and pollen de- reaches 14–22%, while even in mixed beech – position in a mixed deciduous forest II. Seasonal broad-leaved forests at lower levels the beech and annual pollen deposition 1967–1972. Grana, pollen content is much higher. 14: 57–77. 7. In the Tatra Mountains the maximum ARAP R. YA. 1984. Palinologichni doslidzheniya sub- content of European beech pollen is recorded fosilnikh prob z Ukrains’kikh Karpat (summary: in the middle part of beech forest belt, where Palynological analysis of subfossil assays from the Ukrainian Carpathians). Ukrain. Botan. the beech amount in the pollen spectra does Zhurn., 41(l): 73–77. (in Ukrainian). not exceed 26–35% of the total amount of ar- BORATYN´ SKA K. & BORATYN´ SKI A. 1990. Systema- boreal pollen. In the marginal areas and espe- tyka i geograficzne rozmieszczenie (summary: cially at the upper limit of beech forests this Systematics and geographic distribution): 27–75. index drops to 2–5%. There as in the Caucasus In: BIAŁOBOK S. (ed.) Buk zwyczajny – Fagus sylvatica L. PWN, Warszawa-Poznan´ . BRAUN- the main criterion according to which a similar BLANQUET J. 1951. Pflanzensoziologie. Grund- fossil spectrum can be assigned to the beech züge der Vegetationskunde. Springer Verl., Wien. forest spectrum is the predominance of beech CHOCHIEVA K.I. 1980. Uzunlarskaya flora Tskalt- pollen in the group of broad-leaved species. sminda (The Uzunlarian flora of Tskaltsminda). 8. A particularly complex wind regime in Metsniereba, Tbilisi. (in Russian). the Tatra Mountains causes strong averaging DOLUKHANOV A.G. 1941. Kratky ocherk rasti- of the pollen spectra of beech forests, where re- tel’nosti Lagodekhskovo zapovednika (A brief de- gional vegetation features are fixed and better scription of the Lagodekhi Reservation): 11–23. ln: Zaitsev F.A. (ed.) Lagodekhsky zapovednik ( reflected, while in the Caucasus the spectra The Lagodekhi Reservation) lssue 1. lzdat. Akad. better reflect basic features of local vegetation. Nauk Gruzinskoy SSR, Tbilisi. (in Russian). 91

DOLUKHANOV A.G. 1989. Lesnaya rastitel’nost’ Gru- spektrov dlya vosstanovleniya istorii razvitiya go- zii, I (Forest vegetation of Georgia, Part 1). Met- lotsenovoy rastitel’nosti Kolkhidy (summary: On sniereba, Tbilisi. (in Russian). the role of subrecent spore-pollen spectra in the DZWONKO Z. 1990. Ekologia: 237–328. In: BIAŁO- reconstruction of the history of the Holocene BOK S. (ed.) Buk zwyczajny. – Fagus sylvatica L. vegetation in Colchis), lzvest. Akad. Nauk Gru- PWN, Warszawa-Poznan´. (in Polish). zinskoi SSR, Ser.Biolog. 4(4): 250–257. ERDTMAN G. 1943. An introduction to pollen ana- KVAVADZE E.V. & STUCHLIK L. 1990a. Sporovo- lysis. Chronica Botanica, Waltham, Massachu- pyl’tsevye spektry poverkhnostnykh prob iz shiro- setts. kolistvennykh lesov Kintrishskovo i Tsiskariysko- vo zapovednikov – Adzhariya (summary: ERDTMAN G. 1960. The acetolysis method. Svensk. Sporo-pollen spektra of the surface samples from Botan. Tidskr., 54(4): 561–564. the broad-leaved forests of the Kintrishian and GRICHUK V.P. & ZAKLINSKAYA E.D. 1948. Analiz Tsiskarian Forest Reserves, Adzharia). Bull.Acad. iskopaemykh pyl’tsy i spor i evo primenenie v Sci. Georgian SSR, l37(2): 425–428. paleogeografii (Analysis of fossil pollen and KVAVADZE E.V. & STUCHLIK L. 1990b. Subrecent spores and its application in palaeogeography). spore-pollen spectra and their relation to recent Geographizd, Moskva. (in Russian). vegetation belts in Abkhazia (North-Western HESS M. 1965. Pie˛tra klimatyczne w Polskich Karpa- Georgia, USSR). Acta Palaeobot., 30(2): 227–257. tach Zachodnich. Nakładem Uniwersytetu Jagiel- KVAVADZE E. & STUCHLIK L. 1991. Correlation of lon´ skiego, Kraków. (in Polish). subfossil pollen spectra with recent vegetation of KLOPOTOVSKAYA N.B. 1973. Osnovnye zakonomer- the eastern border of the Trialeti Range (The nosti formirovanya sporovo-pyl’tsevykh spektrov Tbilisi environs). Acta Palaeobot., 31(l, 2): v gornykh rayonakh Kavkaza (Basic regularities 273–288. of spore-pollen spectra formation in the Cauca- KVAVADZE E.V. & STUCHLIK L. 1996. Recent pollen sian uplands). Metsniereba, Tbilisi. (in Russian). spectra of the mountain forests of the Lagodekhi KOLAKOVSKY A.A. 1964. Pliotsenovaya flora Kodori Reservation (East Georgia). Acta Palaeobot., (The Pliocene flora of Kodori). lzdat. Akad. Nauk 36(1): 121–147. Gruzinskoy SSR, Sukhumi. (in Russian). LOMINADZE V.P. & CHIRAKADZE G.L. (ed.). 1971. KVAVADZE E.V. 1974. Dannye rezul’tatov sporovo- Klimat i klimaticheskie resursy Gruzii (Climate pyl’tsevovo analiza donnykh otlozheniy reki and climatic resources of Georgia). Gidrometeoiz- Khobi-Zapadnay Gruzya (Data of the spore-pollen dat Leningrad. (in Russian). analysis of bottom sediments of the river Khobi). MAMATSASHVILI N.S. 1972. Sporovo-pyl’tsevye Deponirovanie VINITI (Vsevoiuznii Institut spektry poverkhnostnykh otlozheny doliny reki Nauchnoi i Tekhnicheskoi Informatsii). N 1810- Inguri (Spore-pollen spectra of surface samples of 74, 10 p. (in Russian). the Ingur river). Soobscheniya Akademii Nauk KVAVADZE E.V. 1988. Soderzhanie privnosnoi pyl’tsy Gruzinskoi SSR, 68, 1: 105–108. (in Russian). drevesnykh v sporovopyl’tsevykh spektrakh Kav- MEDWECKA-KORNAs´ A. & KORNAS´ J. 1963. Mapa kaza i Karpat (summary: The content of trans- ros´linnos´ci Ojcowskiego Parku Narodowego (sum- ported arboreal pollen in spore-pollen spectra of mary: Vegetation map of the Ojców National the Caucasus and Carpathians). Bull. Acad. Sc. Park). Ochr. Przyr. 28: 16–87. Georg. SSR, 132(1): 193–196. MEDWECKA-KORNAS´ A. & KORNAS´ J. 1992. The KVAVADZE E.V. 1993. On the interpretation of subfos- Ojców National Park (S. Poland). ln: Veröffentli- sil spore-pollen spectra in the mountains. Acta chungen des Geobotanischen Institutes der Palaeobot., 33(l): 347–360. Eidg.Tech-Hochschule, Stiftung Rübel,in Zürich, KVAVADZE E.V. 1999. The first results of the Pollen 107: 60–82. Monitoring Programme in the Caucasus moun- MICHALIK S. 1978. Ros´liny naczyniowe Ojcowskiego tains (Georgia). Acta Palaeobot., 39(1): 171–177. Parku Narodowego (summary: Vascular plants of KVAVADZE E.V. 2000a. The results of the investiga- the Ojców Natioanl Park). Studia Naturae, A, 16: tion carried out under Pollen Monitoring Pro- 1–171. gramme in the Caucasus mountains on the terri- OBIDOWICZ A. 1996. A late Glacial-Holocene history tory of Georgia. Abstracts of 3-rd Meeting of of the formation of vegetation belts in the Tatra EPMP (12–17 April 2000), National Museum Mts. Acta Palaeobot., 36(2): 159–206. & Gallery, Cardiff, Great Britain. PIDEK A. & BAŁAGA K. 2000. Pollen rain deposition KVAVADZE E.V. 2000b. The role of pollen transport by in the region of Roztocze National Park (SE Po- wind in the formation of pollen spectra from the land) – preliminary results. Abstracts of 3-rd Ukrainian Carpathians. Acta Palaeobot., 40(2): Meeting EPMR (12–17 April 2000), National Mu- 207–214. seum & Gallery, Cardiff, Great Britain. KVAVADZE E. 2001. Annual modern pollen deposition POP E. 1967. Some aeropalynological remarks and con- in the foothills of the Lagodekhi Reservation clusions on the South Carpathian Mountains (Ro- (Caucasus, East Georgia), related to vegetation mania). Rev. Palaeobot. Palynol., 4(1–4): 233–242. and climate. Acta Palaeobot., 41(2): 355–364. RALSKA-JARIEWICZOWA M. 1983. Isopolen maps of KVAVADZE E.V. & STUCHLIK L. 1988. Znachenie Poland: 0–11 000 years B.P. New Phytol., 94: izuchenya subretsentnykh sporovo-pyl’tsevykh 133–175. 92

RUDOLPH K. & FIRBAS F. 1926. Pollenanalitische STOYKO S.N. & ODINAK YA.P. 1988. Bukovye lesa Untersuchung subalpiner Moore des Riesengebir- (Beech forests): 72–77. In: Golubets M.A. (ed.) ges. Ber. Deutsch. Botan. Gesell., 44: 227–238. Ukrainskie Karpaty. Priroda (Ukrainian Carpa- SCHNEIDER R. 1984. Vergleich des Pollengehaltes thians. Nature). Naukova Dumka, Kiev. von Oberflächenproben mit der rezenten Vegeta- STUCHLIK L. & KVAVADZE E.V. 1987. Subrecent tion in Aspromonte, Kalabrien Italien. Disserta- spore-pollen spectra and their relation to recent tiones Batanicae, 72 (Festschrift Welten): 275–318. forest vegetation of Colchis (Western Georgia, SHATILOVA I. I. 1974. Palinologicheskoe obosnovanie USSR). Palaeontographica B, 207(l-6): 133–151. geokhronologii verkhnevo pliotsena i pleistotsena STUCHLIK L. & KVAVADZE E.V. 1995. On the prob- Zapadnoi Gruzii (summary: The palynological lem of actuopalynology in the Carpathians and base of the geochronology of the Upper Pliocene Caucasus. Acta Palaeobot., 35(1): 73–83. and Pleistocene of Western Georgia). Metsniere- S´ RODON´ A. 1985. Fagus in the forest history of Po- ba, Tbilisi. land. Acta Palaeobot., 25(1–2): 119–137. SHATILOVA I.I. 1977. Ob istorii formirovaniya bu- S´ RODON´ A. 1990. Buk w historii lasów Polski (sum- kovykh lesov Kolkhidy (On the history of the Col- mary: beech in the forest history of Poland): 7–27. chis beech forest formation). Isvest. Akad. Nauk In: Bialobok S. (ed.) Buk zwyczajny – Fagus syl- Gruzinskoy SSR, ser. biol., 3, N 5: 458–464. (in vatica L. PWN, Warszawa-Poznan´. Russian). TAKHTADJAN A. 1981. Flowering plants origin and SHATILOVA 1. 1. & RAMISHVILI I.Sh. 1990. Materi- dispersal. Oliver and Boyd, Edinburg. aly po istorii flory i rastitel’nosti Gruzii (Materi- als on the history of flora and vegetation of Geor- TRAMPLER T., KLICZKOWSKA A., DMYTERKO E. gia). Metsniereba, Tbilisi. & SIERIPIN´ SKA A. 1990. Regionalizacja przyrod- niczo-les´na na podstawach ekologiczno-fizjogra- SOKOLOV S. YA. 1951 (ed.). Fagus-buk. In: Derev’ya ficznych (Natural-forestry regionalisation of Po- i kustarniki SSSR, 2, lzdat. Akad. Nauk SSSR, land on the ecological-physiographical basis). Moskva. PWRiL, Warszawa. (in Polish). SOKOLOV S.Ya., SVIAZEVA O.A. & KUBLI V.A. TYUREMNOV S.N. & BEREZINA N.A. 1965. O raz- 1977. Areali derev’ev i kustarnikov SSSR tom 1 rushenii pyl’tsy drevesnikh porod v razlichnikh (Areal of trees and shrubs of the USSR) vol. 1. usloviakh vodno-mineral’novo rezhima (On pollen Nauka, Leningrad. (in Russian). destruction of various tree genera in different STEFANOVA I. 1996. Relationship between recent water-mineral conditions). Vest. Moskov. Gosu- pollen deposition and vegetation in Northern darst. Univer. Ser. Biolog.-Pochvennaya, 5: 62–71. Pirin Mountains. Phytologia Balcanica, 2/2: 61–65. (in Russian).